Ulmus laevis | |
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habitus of an old specimen, Germany | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Rosids |
Order: | Rosales |
Family: | Ulmaceae |
Genus: | Ulmus |
Subgenus: | U. subg. Oreoptelea |
Section: | U. sect. Blepharocarpus |
Species: | U. laevis |
Binomial name | |
Ulmus laevis | |
Distribution map | |
Synonyms | |
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Ulmus laevisPall., variously known as the European white elm, [2] fluttering elm, spreading elm, stately elm and, in the United States, the Russian elm, is a large deciduous tree native to Europe, from France [3] northeast to southern Finland, east beyond the Urals into Kyrgyzstan and Kazakhstan, and southeast to Bulgaria and the Crimea; there are also disjunct populations in the Caucasus and Spain, the latter now considered a relict population rather than an introduction by man, and possibly the origin of the European population. [4] U. laevis is rare in the UK, although its random distribution, together with the absence of any record of its introduction, has led at least one British authority to consider it native. [5] NB: The epithet 'white' elm commonly used by British foresters alluded to the timber of the wych elm. [6]
The species was first identified, as Ulmus laevis, by Pallas, in his Flora Rossica published in 1784. [7] The tree is allogamous and is most closely related to the American elm U. americana . [8]
Endemic to alluvial forest, U. laevis is rarely encountered at elevations above 400 m. [9] Most commonly found along rivers such as the Volga and Danube, it is one of very few elms tolerant of prolonged waterlogged, anoxic ground conditions. The species is threatened by habitat destruction and disturbance in some countries, notably Spain. Flood control schemes are particularly harmful, as seed dispersion is reliant on floods, while abstraction from aquifers lowering ground water levels has compromised the development of the trees.
Although not possessed of an innate genetic resistance to Dutch elm disease, the species is rarely infected in western Europe. [8]
Ulmus laevis is similar in stature to the wych elm, if rather less symmetric, with a looser, untidy, branch structure and less neatly rounded crown. The tree typically reaches a height and breadth of > 30 m, with a trunk < 2 m d.b.h. The extensive shallow root system ultimately forms distinctive high buttresses around the base of the trunk. The bark is smooth at first, then in early maturity breaks into thin grey scales, which separate with age into a network of grey-brown scales and reddish-brown underbark, and finally is deeply fissured in old age like other elms. [10] The leaves are deciduous, alternate, simple ovate with a markedly asymmetric base, < 10 cm long and < 7 cm broad, comparatively thin, often almost papery in texture and very translucent, smooth above with a downy underside. Significantly, the leaf veins do not divide from the central vein to the leaf margin. [11] The leaves are shed earlier in autumn than other species of European elm.
The tree is most reliably distinguished from other European elms by its long flower stems, averaging 20 mm. Moreover, the apetalous wind-pollinated flowers are distinctively cream-coloured, [12] appearing before the leaves in early spring in clusters of 15-30; they are 3–4 mm across. The fruit is a winged samara < 15 mm long by 10 mm broad with a ciliate margin, the single round 5 mm seed maturing in late spring. The seeds have a generally high rate of germination, 45–60% for Serbian trees examined by Stilinović. [13]
Although the species is protandrous, levels of self-pollination can be high [14] The tree can grow very rapidly; where planted in persistently moist soil, trunk width of 13-year-old trees increased by 4 cm per annum at breast height (d.b.h.). [15] The species differs from its closest relative, the American elm, mainly in the irregular crown structure and frequent epicormic shoots, features which also give the tree a distinctive winter silhouette. [16] [17] The American elm also has less acute leaf buds, longer petioles, narrower leaves, and a deeper apical notch in the samara which reaches the seed. [18]
Like other European elms, natural populations of the European white elm have little innate resistance to Dutch elm disease. In a study in France, losses to DED amounted to 28% over a 10 year period. [19] However, research by Irstea has isolated clones able to survive injection with the causal fungus, initially losing < 70% of their foliage, but regenerating strongly the following year. [20]
The tree is not favoured by the vector bark beetles, which colonize it only when there are no other elm alternatives available, [21] an uncommon situation in western Europe. Indeed, in a study of elm in Flanders, not one example of U. laevis was found to be afflicted by Dutch elm disease. [22] Research in Spain has indicated that it is the presence of an antifeedant triterpene, alnulin, at a concentration of 200 μg/g {dried bark} which renders the tree unattractive to the beetles. [23] Ergo: the tree's decline in western Europe has been chiefly owing to woodland clearance in river valleys, and river management systems eliminating flooding, not disease. However, in 2020, it was noted by the Dutch forestry commission that many laevis, but only in Zeeland, were succumbing to Dutch elm disease for reasons unclear. [24]
It was noted by Jouin at Metz, [16] and a century later by Mittempergher and Santini in Italy, that U. laevis had a very low susceptibility to the elm leaf beetle Xanthogaleruca luteola . [25] Research in Germany has established that the tree is also eschewed by the Zig Zag sawfly Aproceros leucopoda . [26]
Elwes observed that trees planted at Ugbrooke in Devon were infested with Cacopsylla ulmi , [27] which he had never found on any other elm in Britain, an affliction confirmed many years later by Richens, who discovered the specimens of U. laevis grown at Kew were the only elms in the Gardens afflicted by the louse, and the aphid Tinocallis platani . [28]
The species has a slight to moderate susceptibility to elm yellows. [25]
U. laevis is essentially a riparian tree, able to withstand over 100 days of continual flooding, [29] although it is intolerant of saline conditions Spanish trees were found to be calcifuge, preferring slightly acid, siliceous soils, and also drought-intolerant, their xylem vessels prone to drought-stress cavitation. [30] In England, the tree failed to prosper in chalk stream valleys, where the soil was predominantly black peat, named 'Adventurers' for the Adventurers' Land SSSI in Cambridgeshire, owing to dehydration in summer. [15] Trees planted in dry ground are notoriously short-lived. [31]
U. laevis is comparatively weak-wooded, much more so than field elm Ulmus minor , and thus an inappropriate choice for exposed locations. In trials in southern England by Butterfly Conservation, young trees of <5 m height were badly damaged by wind gusts of 40 knots (75 km/h) in midsummer storms. [15] The species was never widely introduced to the United States, but is represented at several arboreta. Ulmus effusa, supplied by the Späth nursery of Berlin, was planted at the Dominion Arboretum, Ottawa, Canada, in 1896, [32] as U. pedunculata. [33] In the Far East, the tree has been planted in Xinjiang province and elsewhere in northern China; planting in Tongliao City is known to have been particularly successful. White elm is also known to have been introduced to Australia. [34]
Since the beginning of the 21st century, the tree has enjoyed a small renaissance in England. A popular larval host plant of the white-letter hairstreak Satyrium w-album butterfly across Europe, the elm is now being planted by Butterfly Conservation and other groups to restore local populations decimated by the effects of Dutch elm disease on native or archaeophytic elms. The Cheshire Wildlife Trust, for example, planted numerous white elms on its reserves in the former Vale Royal district of the county. [15]
U. laevis is probably not native to the United Kingdom despite its random occurrence in the countryside, although the date and circumstances of its introduction have not survived. The earliest published references to the tree (as U. effusa, citing Willdenow) were in Sibthorp's Flora Oxoniensis (1794), [35] and (as U. effusaWilld. but without description) in Miller's posthumously revised Gardener's and Botanist's Dictionary (1807). [36] The first specimen to be reported in cultivation, in 1838, was at Whiteknights Park, Reading, which featured an elm grove; [37] the tree measured 63 ft (19 m) in height, suggesting it had been planted at the end of the 18th century. [38] However, the authenticity of the Whiteknights tree is a matter of contention; it flowered but did not set fertile seed, which suggested to Loudon that it might be U. campestris (U. minor 'Atinia'), or, on account of it not producing suckers, possibly U. montana (:U. glabra). [38] Moreover, Whiteknights was supplied by the Lee and Kennedy nursery of Hammersmith, [39] which is not known to have stocked U. laevis. A tree at Syon Park identified by Elwes & Henry as U. laevis was later considered by Bean as more closely resembling U. americana by dint of its symmetrical branch arching. [17] The species was not reported from the wild until 1943, with the discovery of a tree in a Surrey hedgerow. [40]
It is possible the tree's distribution was associated with Capability Brown (1716–1783), known to have favoured U. laevis, which he listed among his preferred "native" (sic) trees. [41] [ non-tertiary source needed ] This could explain the existence of the seven old specimens discovered by Elwes in 1908 on Mount Pleasant within Ugbrooke Park, Devon, [16] designed by Brown in 1761. [42] Ugbrooke is four miles from Mamhead Park, which had earlier been planted with numerous exotic trees, notably holm oak, collected by its owner, merchant Thomas Ball ( d. 1749) during his commercial travels in Europe. [43] [44] Ball's introductions were known to have been marketed by his head gardener William Lucombe, who in 1720 founded the first commercial nursery in the south-west at Exeter, [45] though an account of trees growing at Mamhead by Pince (grandson of Lucombe) in 1835 makes no mention of U. laevis nor of any other elms. [46] None of Lucombe's early catalogues are known to survive, and thus the introduction of U. laevis through south Devon cannot be confirmed. However, the tree does not feature in any of the surviving arboreta accessions lists, or catalogues of the larger, nationally famous, nurseries of the day, and its earliest-known mention in commerce remains in the south-west, in the catalogue of the Ford & Please nursery (as U. pedunculata) at Exeter circa 1836. [45] James Main mentions the tree as 'a native of Hungary' and in 1838 only to be met in 'ornamental plantations', [47] but by 1846 was 'becoming available in (UK) nurseries'. [48]
U. laevis, obtained from the Späth nursery of Berlin as U. effusa, was planted in Kew Gardens (1895), [16] in the Ryston Hall arboretum, Norfolk (1914), [49] [50] and, re-propagated, in Cambridge University Botanic Garden (1909). Evidently the tree did not gain in popularity, and was overlooked or ignored by most authors of popular guides to trees in Britain during the 20th century, notably Mabey in his Flora Britannica. [51] The tree is also omitted from Keble-Martin's comprehensive Flora of Devon. [52]
It is not known whether U. laevis was introduced to Scotland before the early 20th century. Two of the three specimens supplied by the Späth nursery, Berlin, to the Royal Botanic Garden Edinburgh in 1902 as U. effusa may survive in Edinburgh, as it was the practice of the garden to distribute trees about the city; [53] the third specimen was in the garden itself. [54] Other examples can be found in the city, notably in Fettes College grounds opposite Inverleith Allotments, and at the entrance to North Merchiston Cemetery. [55] In Wales, two mature trees with numerous seedlings occur in a small wood at Rhydyfelin near Aberystwyth, [56] while another grows at Llandegfan, Anglesey. In Ireland, the tree is represented by a line of four at the Old Rectory, Kells Road, Ardee, County Louth (22 m or 72 ft, girth 3 m or 9.8 ft, October 2009), [57] and in the Channel Islands, by a clump near the well at La Seigneurie (Le Manoir), Sark. [58]
The two largest known trees in Europe are at Gülitz in Germany (3.1 m d.b.h.), and at Komorów in Poland (2.96 m d.b.h. in 2011), known as the Witcher. Other veterans survive at Casteau, Belgium (bole-girth 5.15 m), in Rahnsdorf near Berlin (bole-girth 4.5 m) [60] [61] and in Ritvala, Finland (bole-girth 4.49 m). [62] A lane of Ulmus laevis is found at Eibergen, Netherlands (see Gallery below), while a large, mature specimen is found within the Alhambra, Granada.
Ulmus laevis has very occasionally been planted as an ornamental tree in the UK, and even more randomly in countryside hedgerows. The UK Champion is at Ferry Farm, on the banks of the Tamar at Harewood, Cornwall (27 m high, 1.8 m d.b.h. in 1997). [63] Other examples are few and far between though sometimes of considerable age, surviving amid diseased native elm in Cornwall at Torpoint, [64] and Pencalenick (21 m high, d.b.h. 1.75 m), [65] and near Over Wallop in Hampshire (16 m high, d.b.h. 1.3 m 2016) [66] The largest-known aggregation in England is the ring of 50 trees planted circa 1950 within a ring of common lime around a former ammunition dump on the elevated chalk of Salisbury Plain at Hexagon Wood, Larkhill, about 2 km (1.2 mi) north of Stonehenge. [15]
In the United States, a tree of 31.4 m (103 ft) in height (2015) grows at 3331 NE Hancock Street in Portland, Oregon; its age is not known. [67] [68]
The timber of the white elm is of poor quality, the cross-grain causing problems when machined, and thus of little practical use, not even as firewood.[ citation needed ] The density of the timber is significantly lower than that of other European elms. However, owing to its rapid growth, tolerance of soil compaction, air pollution and de-icing salts, the tree has long been used for amenity planting in towns and along roadsides. [8]
U. laevis is easily grown from seed sown on ordinary compost and kept well-watered. However, viability can vary greatly from year to year, while the seed is remarkably short-lived. Germination should occur within one week even without heat, the best seedlings attaining as much as half a metre in their first year. [69] Softwood cuttings taken in June is also a reliable method; the cuttings strike very quickly, well within a fortnight, rapidly producing a dense matrix of roots.
Several putative varieties have been identified. A var. celtidea from Ukraine was reported by Rogowicz in the middle of the 19th century, but no examples are known to survive. Another, var. parvifolia, has been reported from Serbia. [70] A third, var. simplicidens, is very rare; the only example known to survive is at the National Botanic Garden of Latvia in Salaspils. Kew had a grafted var. glabra in the early 20th century (provenance unknown), [71] [72] a clone of which is present at Wakehurst Place. [73]
Compared with the other European species of elm, U. laevis has received scant horticultural attention, there being only eight recorded cultivars:
In Russia other ornamental forms are recognized: f. argentovariegata, f. rubra, and f. tiliifolia. A pyramidal form was reported in 1888 from the Fredericksfelde cemetery in Berlin by Bolle. [74] A line of similar monopodial trees grows (2019) on the island in the Lot at Entraygues, France.
U. laevis does not hybridize naturally, in common with the American elm (U. americana) to which it is closely related. However, in experiments at the Arnold Arboretum, it was successfully crossed with U. thomasii and U. pumila; no such crosses have ever been released to commerce.
Ulmus davidiana var. japonica, the Japanese elm, is one of the larger and more graceful Asiatic elms, endemic to much of continental northeast Asia and Japan, where it grows in swamp forest on young alluvial soils, although much of this habitat has now been lost to intensive rice cultivation.
Ulmus × hollandica 'Vegeta', sometimes known as the Huntingdon Elm, is an old English hybrid cultivar raised at Brampton, near Huntingdon, by nurserymen Wood & Ingram in 1746, allegedly from seed collected at nearby Hinchingbrooke Park. In Augustine Henry's day, in the later 19th century, the elms in Hinchingbrooke Park were U. nitens. Richens, noting that wych elm is rare in Huntingdonshire, normally flowering four to six weeks later than field elm, pointed out that unusually favourable circumstances would have had to coincide to produce such seed: "It is possible that, some time in the eighteenth century, the threefold requirements of synchronous flowering of the two species, a south-west wind", "and a mild spring permitting the ripening of the samaras, were met."
Ulmus bergmannianaC.K.Schneid., commonly known as Bergmann's elm, is a deciduous tree found across much of China in forests at elevations of 1500–3000 m.
Ulmus castaneifoliaHemsley, the chestnut-leafed elm or multinerved elm, is a small deciduous tree found across much of China in broadleaved forests at elevations of 500–1,600 metres (1,600–5,200 ft).
Ulmus laciniata(Trautv.) Mayr, known variously as the Manchurian, cut-leaf, or lobed elm, is a deciduous tree native to the humid ravine forests of Japan, Korea, northern China, eastern Siberia and Sakhalin, growing alongside Cercidiphyllum japonicum, Aesculus turbinata, and Pterocarya rhoifolia, at elevations of 700–2200 m, though sometimes lower in more northern latitudes, notably in Hokkaido.
Ulmus macrocarpaHance, the large-fruited elm, is a deciduous tree or large shrub endemic to the Far East excluding Japan. It is notable for its tolerance of drought and extreme cold and is the predominant vegetation on the dunes of the Khorchin sandy lands in the Jilin province of north-eastern China, making a small tree at the base of the dunes, and a shrub at the top.
Ulmus chenmouiW. C. Cheng, commonly known as the Chenmou, or Langya Mountain elm, is a small deciduous tree from the more temperate provinces of Anhui and Jiangsu in eastern China, where it is found at elevations below 200 m on the Langya Shan and Baohua Shan mountains. The tree was unknown in the West until 1979, when seeds were sent from Beijing to the De Dorschkamp research institute at Wageningen in the Netherlands.
Ulmus davidiana, also known as the David elm, or Father David elm, is a small deciduous tree widely distributed across China, Mongolia, Korea, Siberia, and Japan, where it is found in wetlands along streams at elevations of 2000–2300 m (6,500–7,500 ft). The tree was first described in 1873 from the hills north of Beijing, China.
Ulmus villosa, the cherry-bark elm or Marn elm, is one of the more distinctive Asiatic elms, and a species capable of remarkable longevity. It is endemic to the valleys of the Kashmir at altitudes of 1,200–2,500 metres but has become increasingly rare owing to its popularity as cattle fodder. Mature trees are now largely restricted to temples and shrines where they are treated as sacred. Some of these trees are believed to be over 800 years old.
The field elm cultivar Ulmus minor 'Stricta', known as Cornish elm, was commonly found in South West England, Brittany, and south-west Ireland, until the arrival of Dutch elm disease in the late 1960s. The origin of Cornish elm in England remains a matter of contention. It is commonly assumed to have been introduced from Brittany. It is also considered possible that the tree may have survived the ice ages on lands to the south of Cornwall long since lost to the sea. Henry thought it "probably native in the south of Ireland". Dr Max Coleman of Royal Botanic Garden Edinburgh, arguing in his 2002 paper on British elms that there was no clear distinction between species and subspecies, suggested that known or suspected clones of Ulmus minor, once cultivated and named, should be treated as cultivars, preferred the designation U. minor 'Stricta' to Ulmus minor var. stricta. The DNA of 'Stricta' has been investigated and the cultivar is now known to be a clone.
The Field Elm cultivar Ulmus minor 'Sarniensis', known variously as Guernsey elm, Jersey elm, Wheatley elm, or Southampton elm, was first described by MacCulloch in 1815 from trees on Guernsey, and was planted in the Royal Horticultural Society's gardens in the 1820s. It was listed in the Loddiges catalogue of 1836 as Ulmus sarniensis and by Loudon in Hortus lignosus londinensis (1838) as U. campestris var. sarniensis. The origin of the tree remains obscure; Richens believed it "a mutant of a French population of Field elm", noting that "elms of similar leaf-form occur in Cotentin and in northern Brittany. They vary much in habit but some have a tendency to pyramidal growth. Whether the distinctive habit first developed on the mainland or in Guernsey is uncertain."
The hybrid elm cultivar Ulmus × hollandica 'Belgica', one of a number of hybrids arising from the crossing of Wych Elm with a variety of Field Elm, was reputedly raised in the nurseries of the Abbey of the Dunes, Veurne, in 1694. Popular throughout Belgium and the Netherlands in the 19th century both as an ornamental and as a shelter-belt tree, it was the 'Hollandse iep' in these countries, as distinct from the tree known as 'Dutch Elm' in Great Britain and Ireland since the 17th century: Ulmus × hollandica 'Major'. In Francophone Belgium it was known as orme gras de Malines.
The hybrid elm cultivar Ulmus × hollandica 'Dampieri', one of a number of cultivars arising from the crossing of the Wych Elm U. glabra with a variety of Field Elm U. minor, is believed to have originated in continental Europe. It was marketed in Wetteren, Belgium, in 1851 as 'Orme de Dampier', then in the Low Countries in 1853, and later identified as Ulmus campestris var. nuda subvar. fastigiata DampieriHort., Vilv. by Wesmael (1862).
The hybrid elm cultivar Ulmus × hollandica 'Wredei', also known as Ulmus × hollandica 'Dampieri Aurea' and sometimes marketed as Golden Elm, originated as a sport of the cultivar 'Dampieri' at the Alt-Geltow Arboretum, near Potsdam, Germany, in 1875.
Ulmus 'Exoniensis', the Exeter elm, was discovered near Exeter, England, in 1826, and propagated by the Ford & Please nursery in that city. Traditionally believed to be a cultivar of the Wych Elm U. glabra, its fastigiate shape when young, upward-curving tracery, small samarae and leaves, late leaf-flush and late leaf-fall, taken with its south-west England provenance, suggest a link with the Cornish Elm, which shares these characteristics. The seed, however, is on the stalk side of the samara, a feature of wych elm and its cultivars, whereas in hybrids it would be displaced towards the notch.
The Wych Elm cultivar Ulmus glabra 'Horizontalis', commonly known as the Weeping Wych Elm or Horizontal Elm, was discovered in a Perth nursery circa 1816. The tree was originally identified as 'Pendula' by Loddiges (London), in his catalogue of 1836, a name adopted by Loudon two years later in Arboretum et Fruticetum Britannicum, 3: 1398, 1838, but later sunk as a synonym for 'Horizontalis'.
The Wych Elm cultivar Ulmus glabra 'Cornuta', in cultivation before 1845 – Fontaine (1968) gives its provenance as France, 1835 – is a little-known tree, finally identified as a cultivar of U. glabra by Boom in Nederlandse Dendrologie 1: 157, 1959.
Ulmus laevis var. celtideaRogow. [: like Celtis, the leaves] is a putative variety of European White Elm first described by Rogowicz, who found the tree in 1856 along the river Dnjepr near Chernihiv in what is now northern Ukraine. The type specimen is held at the National Herbarium of Ukraine. The variety was first named as Ulmus pedunculata var. celtidea. Litvinov (1908) considered it a species, calling it Ulmus celtideaLitv., a view not upheld by other authorities.
The 'dwarf' elm cultivar Ulmus 'Jacqueline Hillier' ('JH') is an elm of uncertain origin. It was cloned from a specimen found in a private garden in Selly Park, Birmingham, England, in 1966. The garden's owner told Hillier that it might have been introduced from outside the country by a relative. Hillier at first conjectured U. minor, as did Heybroek (2009). Identical-looking elm cultivars in Russia are labelled forms of Siberian Elm, Ulmus pumila, which is known to produce 'JH'-type long shoots. Melville considered 'JH' a hybrid cultivar from the 'Elegantissima' group of Ulmus × hollandica. Uncertainty about its parentage has led most nurserymen to list the tree simply as Ulmus 'Jacqueline Hillier'. 'JH' is not known to produce flowers and samarae, or root suckers.
The Siberian elm cultivar Ulmus pumila 'Pinnato-ramosa' was raised by Georg Dieck, as Ulmus pinnato-ramosa, at the National Arboretum, Zöschen, Germany, from seed collected for him circa 1890 in the Ili valley, Turkestan by the lawyer and amateur naturalist Vladislav E. Niedzwiecki while in exile there. Litvinov (1908) treated it as a variety of Siberian elm, U. pumilavar.arborea but this taxon was ultimately rejected by Green, who sank the tree as a cultivar: "in modern terms, it does not warrant recognition at this rank but is a variant of U. pumila maintained and known only in cultivation, and therefore best treated as a cultivar". Herbarium specimens confirm that trees in cultivation in the 20th century as U. pumilaL. var. arboreaLitv. were no different from 'Pinnato-ramosa'.
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