Ilex aquifolium

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Ilex aquifolium
Ilex-aquifolium (Europaeische Stechpalme-1).jpg
Leaves and fruit
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Aquifoliales
Family: Aquifoliaceae
Genus: Ilex
Species:
I. aquifolium
Binomial name
Ilex aquifolium
L.
Ilex aquifolium range.svg
Distribution map of Ilex aquifolium
Ilex aquifolium Ilex aquifolium in the Tashkent Botanical Garden.jpg
Ilex aquifolium

Ilex aquifolium, the holly, common holly, English holly, European holly, or occasionally Christmas holly, is a species of flowering plant in the family Aquifoliaceae, native to western and southern Europe, northwest Africa, and southwest Asia. [2] [3] [4] It is regarded as the type species of the genus Ilex , which by association is also called holly. It is an evergreen tree or shrub found, for example, in shady areas of forests of oak and in beech hedges. In the British Isles it is one of very few native hardwood evergreen trees. It has a great capacity to adapt to different conditions and is a pioneer species that repopulates the margins of forests or clearcuts.

Contents

I. aquifolium can exceed 10 m in height, but is often found at much smaller heights, typically 2–3 m (6.6–9.8 ft) tall and broad, with a straight trunk and pyramidal crown, branching from the base. It grows slowly and does not usually fully mature due to cutting or fire. It can live 500 years, but usually does not reach 100.

Ilex aquifolium is the species of holly long associated with Christmas, and previously the Roman festival of Saturnalia. Its glossy green prickly leaves and bright red berries (produced only by the female plant) are represented in wreaths, garlands and cards wherever Christmas is celebrated. It is a subject of music and folklore, especially in the British tradition. It is also a popular ornamental shrub or hedge, with numerous cultivars in a range of colours.

Description

Frosted foliage and berries Ilex aquifolium berries rime.jpg
Frosted foliage and berries
European holly flowers; male above, female below (leaves cut to show flowers more clearly) Hollyflowers.jpg
European holly flowers; male above, female below (leaves cut to show flowers more clearly)

In exceptional cases European holly can grow to 25 m (82 ft) in height, [5] but more typically is 3 to 15 m (10 to 49 ft) tall and often only grows into a shrub rather than reaching tree stature. [6] It has a woody stem as wide as 40–80 cm (16–31 in), rarely 100 cm (39 in) or more, in diameter. [7] The bark is smooth and silver-grey, but with age will develop small fissures. [6] The root system is a fibrous mat near the surface with only a few roots that descend to greater depths. In dense stands the roots will spread only about as wide as the branches. [8] Holly trees only rarely produce suckers, but those that do can produce a clump stems. Lower branches that become burried in dead leaves can also sometimes root. [9]

The leaves are 5–12 cm long and 2–6 cm broad; [7] they are evergreen, and though they can last for as much as eight years less than five is more typical. [10] In the upper canopy of trees exposed to full sun leaves fall after just one or two years. [11] They are dark and shining green on the upper surface and lighter on the underside, their shape is ovate to oblong-elliptic, broader at the base and rounded like the outline of an egg to somewhat rectangular with curving sides. [5] In the young and in the lower limbs of mature trees, the leaves have three to five sharp spines on each side, pointing alternately upward and downward, while leaves of the upper branches in mature trees lack spines. [7] [4] In rare cases, a tree may lack spines even on its lower leaves. [12]

The flowers are white, four-lobed, and pollinated by bees. Holly is usually dioecious, meaning that flowers that produce fruit and flowers that produce pollen are on separate trees. [10] The sex cannot be determined until the plants begin flowering, usually between 4 and 12 years of age. In male specimens, the flowers are yellowish and appear in axillary groups. In the female, flowers are isolated or in groups of three and are small and white or slightly pink, and consist of four petals and four sepals partially fused at the base. Though spring is the standard time for flowering, individual trees have been known to bloom in winter as late as January. [9]

The fruit only appears on female plants, which require male plants nearby to fertilise them. The fruit is a drupe (stone fruit), about 6–10 mm in diameter, a bright red or bright yellow, which matures around October or November; at this time they are very bitter due to the ilicin content [13] and so are rarely eaten until late winter after frost has made them softer and more palatable. They are eaten by rodents, birds and larger herbivores. Each fruit contains 3 to 4 seeds which typically germinate in the second or third spring. [14]

Taxonomy

Ilex aquifolium was given its scientific name in 1753 by Linnaeus. He placed it in the genus Ilex which is classified in the family Aquifoliaceae. [15] It is also designated as the type for the genus, the species that defines its genus. [16] More than 100 botanical varieties of Ilex aquifolium have been described by botanists, but none of them are accepted names according to Plants of the World Online (POWO). Together with 23 species and other names it has a total of 145 synonyms, though for brevity none of the varieties or forms of I. aquifolium are presented here. [15]

Table of Synonyms
NameYearRankNotes
Aquifolium croceumRaf.1838species= het.
Aquifolium feroxMill. ex Rafin.1838species= het.
Aquifolium heterophyllumRaf.1838species= het.
Aquifolium ilexScop.1771species= het.
Aquifolium lanceolatumRaf.1838species= het.
Aquifolium planifoliumRaf.1838species= het.
Aquifolium spinosumLam.1779species= het.
Aquifolium undulatumRaf.1838species= het.
Aquifolium vulgareSt.-Lag.1880species= het.
Ilex balearicaDesf.1809species= het.
Ilex balearica var. cordataGöpp.1852variety= het.
Ilex chrysocarpaWender.1828species= het.
Ilex ciliataA.Vilm.1860species= het.
Ilex citriocarpaMurr1912species= het.
Ilex crassifoliaAiton ex Steud.1840species= het.
Ilex echinataMill.1768species= het.
Ilex ferox(Aiton) Reider1835species= het.
Ilex fischeriCarrière1887species= het.
Ilex maderensisWilld.1814species= het., nom. illeg.
Ilex myrtifolia var. aureomaculataVan Geert1875variety= het.
Ilex nigricans(Göpp.) A.Henry1913species= het.
Ilex perado subsp. iberica(Loes.) S.Andrews1984subspecies= het.
Ilex perado var. ibericaLoes.1901variety= het.
Ilex platyphyllaBooth1854species= het.
Ilex sempervirensSalisb.1796species= het.
Ilex vulgarisGray1821species≡ hom., nom. superfl.
Notes: ≡ homotypic synonym  ; = heterotypic synonym

Names

The genus Ilex was origionally the Latin name for the holly oak (Quercus ilex). The species name, aquifolium, is Botanical Latin coming from the Latin acus meaning needle and folia meaning leaf. [17]

The name holly can be used to mean any species in the genus Ilex, but originally referred to Ilex aquifolium and continues to be used to mean this species particularly in addition to names such as European holly. Holly is a shortened form of the Old English holegn or holen. [18] In Scotland holly is still sometimes called by the older names hollin or hollen. [19] The Middle English usage of hollin also became a name for a grove of holly trees, especially one that was regularly cut for to feed domestic animals and it is part of some place names in England. [20] It is also known by the common names common holly and English holly. [21] [22] Both Ilex aquifolium and the North American Ilex opaca are occasionally called Christmas holly. [23]

Smooth leaved holly is known as free holly or slike holly in Shropshire. [9] Locally, it has been known as holm, holme, or home in West Devon and Cornwall. [24]

Distribution and habitat

Holly is native to much of the western half of Europe as well as northwestern Africa. [15] Its native status in central and Eastern Europe is disputed. According to Plants of the World Online (POWO) it is not native or reproducing in Poland or Hungary, but is native to Bulgaria and Romania. [15] According to the International Union for Conservation of Nature (IUCN) its status in Romania is uncertain and it is introduced and naturalized to some extant in Hungary, though they agree that it is not found in Poland or eastward in northern Europe. [1]

In Asia its status is disputed with POWO listing it as an introduced species in Turkey and Syria while the IUCN lists it as native and also native to Iran and Lebanon. [15] [1]

The species is shifting northwards in Norway and towards to the northeast in Denmark and Germany. Holly grows from sea level to elevations as high as 600 m (2,000 ft). [1]

Holly is a common species in its range and widespread. Because of this it is a species of least-concern according to the IUCN, however there is a decline in mature individuals and its population is severely fragmented. Though, local populations may face threats. In the country of Sweden only one wild tree remains. [1]

It grows in the undergrowth of oak forest and beech forest in particular, although at times it can form a dense thicket as the dominant species. It requires moist, shady environments, found within forests or in shady slopes, cliffs, and mountain gorges. [2] [7]

Along the west coast of the United States and Canada, from California to British Columbia, [25] non-native English Holly has proved very invasive, quickly spreading into native forest habitat, where it thrives in shade and crowds out native species. It has been placed on the Washington State Noxious Weed Control Board's monitor list, and is a Class C invasive plant in Portland. [26] [27] [28]

During the Cenozoic Era, the Mediterranean region, Europe, and northwest Africa had a wetter climate and were largely covered by laurel forests. Holly was a typical representative species of this biome, where many current species of the genus Ilex were present. With the drying of the Mediterranean Basin during the Pliocene, the laurel forests gradually retreated, replaced by more drought-tolerant sclerophyll plant communities. The modern Ilex aquifolium resulted from this change.[ clarification needed ][ citation needed ] Most of the last remaining laurel forests around the Mediterranean are believed to have died out approximately 10,000 years ago at the end of the Pleistocene.

Ecology

Holly thicket in the West midlands, England. Holly thicket, Holly Hurst, Sutton Park - geograph.org.uk - 1859863.jpg
Holly thicket in the West midlands, England.

Holly is a rugged pioneer species that prefers relatively moist areas, and tolerates frost as well as summer drought. With its spiny leaves it can form a dense thicket that defends other woody species from herbivores aiding the regeneration of forests. [29] The plant is common in the garrigue and maquis and is also found in deciduous forest and oak forest.

Pure stands of hollies can grow into a labyrinth of vaults in which thrushes and deer take refuge, while smaller birds are protected among their spiny leaves. After the first frost of the season, holly fruits become soft and fall to the ground serving as important food in its native regions for winter birds at a time of scarce resources.

The flowers are attractive as nectar sources for insects such as bees, wasps, flies, and small butterflies. The commonly-encountered pale patches on leaves are due to the leaf-mine insect Phytomyza ilicis .

It is an invasive species on the West Coast of Canada [25] and the United States as well as in Hawaii. [28] [30]

Epigenetics

Holly is well known in epigenetics. Some cultivars have smooth leaf edges, or both smooth and prickly leaf edges on the same plant. In response to stress these cultivars can produce leaves with more prickles. [31]

Cultivation

One of several variegated cultivars 2014-08-29 13 46 38 Variegated English Holly at the Pinelands Preservation Alliance headquarters in Southampton Township, New Jersey.JPG
One of several variegated cultivars

Ilex aquifolium is widely grown in parks and gardens in temperate regions. [32] Hollies are often used for hedges; [33] the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping. [34] Holly can grow in a wide variety of difficult soils including clay, acidic, and nutrient poor soils, but is intollerant of poor drainage. In wet conditions it is likely to have problems such as root rot, canker, or leaf blight. [35]

AGM cultivars

Numerous cultivars have been selected, [36] of which the following have gained the Royal Horticultural Society's Award of Garden Merit: [37]

  • I. aquifolium [38]
  • 'Amber' (female) [39]
  • 'Argentea Marginata' [40]
  • 'Ferox Argentea' [41]
  • 'Golden Queen' [42]
  • 'Handsworth New Silver' [43]
  • 'J.C. van Tol' [44]
  • 'Madame Briot' [45]
  • 'Pyramidalis' [46]
  • 'Silver Queen' [47]

Ilex × altaclerensis

The hybrid Ilex × altaclerensis was developed at Highclere Castle in Hampshire, England, in 1835, a cross between I. aquifolium and the tender species I. perado. The following cultivars have gained the RHS AGM:

Chemistry and toxicity

Holly berries contain alkaloids, theobromine, saponins, caffeic acid, and a yellow pigment, ilixanthin. [13] [52] [53] The berries are generally regarded as toxic to humans. [52] [53]

Uses

As early as the 1200s and into the first part of the 1700s Ilex aquifolium was cultivated for use as winter fodder for many domestic animals. Sheep were most frequently fed holly, but horses and cattle were also given it and branches were cut down for red deer. [54] This practice is best documented in southern areas of the Pennines near Sheffield, [54] but might have been more widespread in Britain and there are a few instances of it recorded in northwest France. [55] The smooth leaves were specifically selected to be cut in at least some instances. [56] Though the practice is much rarer, branches are still cut for sheep or cattle in Dumfries, Derbyshire, Cumbria, and in the New Forest. [57] It was also practiced in Ireland until the middle of the twentith century. [33]

The wood of the holly is hard and white with a slight grey or green tinge and without a noticeable heartwood. It is fine grained and quite heavy. It is difficult to dry and is often cut into small pieces avoid problems with it warping. It is used for wood inlay, small fancy carving, or small turned peices. It is easily stained black and is occasionally substituted for ebony. [58] In the 1800s holly was extensively used, and considered the best material, for making driving whips for horse drawn carriages. [59] Although popular stories suggest that holly and other locally produced woods were used to make Great Highland bagpipes the evidence shows that it was always made using imported woods in the lowlands. [60]

Holly, like ash trees, makes a desirable firewood due to being flamable while still green. [61]

The leaves are prepared as a tea, though rarely, as a folk medicine for pain relief in the Pallars region in the mountains of northwestern Catalonia. [62]

In culture

In the British Isles common holly has been used in rituals and as a symbol both before and after the coming of Christianity in similar ways to the use of evergreen plants during midwinter in many cultures. [63] Branches of holly continue to be used in place of palm fronds on Palm Sunday in some areas of Europe. It is also often used in Christmas decorations, particularly in Britain, its evergreen leaves a symbol of life continuing during the winter season. [64] Though there is no traditional prohibition against cutting boughs or twigs, cutting down a holly tree is believed to bring bad luck. [65] Like other spiny plants, the common holly is believed to protect against malicious witchcraft. [66] On the fells in Lancashire branches of holly are placed in new buildings or where an animal has died by the superstitious. [67] A similar pratice in France is to hang a branch of holly above animal pens to keep away ringworm. [59]

The name of the hamlet of Hulver in Suffolk is the same at the Middle English name of the species, though in 1997 there were only 18 holly trees there. There is a similar lack of the namesake trees in Hollybush near the border between Worcestershire and Herefordshire. [9]

References

  1. 1 2 3 4 5 Barstow & Khela 2018.
  2. 1 2 Flora Europaea: Ilex aquifolium
  3. Med-Checklist: Ilex aquifolium
  4. 1 2 Flora of NW Europe: Ilex aquifolium Archived 2009-06-09 at the Wayback Machine
  5. 1 2 Humphries, Press & Sutton 2000, p. 244.
  6. 1 2 Press & Hosking 1992, p. 196.
  7. 1 2 3 4 Rushforth 1999.
  8. Peterken & Lloyd 1967, pp. 850–851.
  9. 1 2 3 4 Mabey 1997, p. 245.
  10. 1 2 Peterken & Lloyd 1967, p. 841.
  11. Peterken & Lloyd 1967, p. 851.
  12. Fitter 1999, p. 200.
  13. 1 2 Morck 1978.
  14. Peterken & Lloyd 1967, p. 854.
  15. 1 2 3 4 5 POWO 2025.
  16. Britton & Brown 1913, p. 486.
  17. Christenhusz, Fay & Chase 2017, p. 589.
  18. OED 2025b.
  19. OED 2025a.
  20. Mabey 1997, p. 246.
  21. Hyde & Harrison 1977, p. 78.
  22. Krüssmann 1986, p. 172.
  23. Galle 1998, p. 539.
  24. Britten & Holland 1886, p. 264.
  25. 1 2 "Ring in the holiday season and reduce the spread of invasive species! -". Invasive Species Council of British Columbia | ISCBC Plants & Animals. 2020-02-10. Retrieved 2020-12-27.
  26. "Washington State Noxious Weed Control Board".
  27. "English Holly | Invasive Species Council of British Columbia | ISCBC Plants & Animals". Archived from the original on 2015-09-06. Retrieved 2015-08-14.
  28. 1 2 "Ilex aquifolium (English holly)". California Invasive Plant Council. Archived from the original on 2016-08-16. Retrieved 2016-10-02.
  29. Kingsbury 2014, p. 66.
  30. "English Holly - Ilex aquifolium". King County, Washington. Archived from the original on 2011-02-27. Retrieved 2011-01-30.
  31. "#bioPGH blog – to Prickle or Not to Prickle: Holly Leaves and Epigenetics | Phipps Conservatory and Botanical Gardens | Pittsburgh PA".
  32. Phillips & Grant 1978, p. 122.
  33. 1 2 Nelson & Walsh 1994, p. 42.
  34. Northumbria Police: Security starts at the Garden Gate Archived 2008-11-08 at the UK Government Web Archive
  35. NCCE Staff 2025.
  36. Brickell & Chisholm 2003, p. 562.
  37. "AGM Plants - Ornamental" (PDF). Royal Horticultural Society. July 2017. p. 43. Retrieved 3 March 2018.
  38. "RHS Plant Selector Ilex aquifolium AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  39. "RHS Plant Selector Ilex aquifolium 'Amber' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  40. "RHS Plant Selector Ilex aquifolium 'Argentea Marginata' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  41. "RHS Plant Selector Ilex aquifolium 'Ferox Argentea' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-10.
  42. "RHS Plant Selector Ilex aquifolium 'Golden Queen' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  43. "RHS Plant Selector Ilex aquifolium 'Handsworth New Silver' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  44. "RHS Plant Selector Ilex aquifolium 'J.C. van Tol' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  45. "RHS Plant Selector Ilex aquifolium 'Madame Briot' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  46. "RHS Plant Selector Ilex aquifolium 'Pyramidalis' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  47. "RHS Plant Selector Ilex aquifolium 'Silver Queen' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  48. "RHS Plant Selector Ilex × altaclerensis 'Belgica Aurea' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  49. "RHS Plant Selector Ilex × altaclerensis 'Camelliifolia' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  50. "RHS Plant Selector Ilex × altaclerensis 'Golden King' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  51. "RHS Plant Selector Ilex × altaclerensis 'Lawsoniana' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  52. 1 2 Leikin & Paloucek 2002, p. 80.
  53. 1 2 Turner, Nancy J.; P. von Aderkas (2009). The North American Guide to Common Poisonous Plants and Mushrooms. Timberpress. p. 210. ISBN   978-0-88192-929-4.
  54. 1 2 Spray 1981, p. 103.
  55. Spray 1981, p. 97.
  56. Spray 1981, p. 99.
  57. Mabey 1997, p. 248.
  58. Bennett et al. 1956, p. 109.
  59. 1 2 Mabey 1997, p. 249.
  60. Cheape 2008, p. 134.
  61. Thomas 2000, p. 250.
  62. Agelet & Vallès 2003, p. 213.
  63. Bailes 2006, p. 24.
  64. Spohn & Spohn 2008, p. 115.
  65. Mabey 1997, pp. 244–245.
  66. Galle 1998, p. 18.
  67. Mabey 1997, pp. 248–249.

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