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Never ignore a holly

  • Green Fingers
  • Mar 30
  • 16 min read

 

The lonely female holly
The lonely female holly

The tree looked lonely, growing from the base of a crag high above the Lakeland town of Ambleside. It was a female holly (Ilex aquifolium), its bright red berries (drupes) waiting for a passing pollinator. Yet there were no other trees, regardless of species, within at least two kilometres. Trees do communicate[i], indeed all plants inter-relate[ii], which made me ponder about hollies. It is a fact that just like a mirror of modern society, trees can suffer from loneliness[iii]. I stopped, checked there was no one nearby to declare me insane, doffed my mountain hat and said, “Good morning.” For a moment, I imagined the lady holly nodded her reply. Just for a few seconds she had received human company. I then walked on.

 

I walk past plenty of hollies in the Lake District. They need a moist but well-drained soil, enriched with plenty of organic matter and will grow in most soils, acidic or alkaline. They steer clear of solid chalk[iv]. Ideally, a holly seeks full sun, although will handle shade. Often alone and independent, hollies can also grow together, in an almost-clump. When I see this, I call it a hollinus, similar to a salicetum, which is a place dominated by willow trees, or quercetum, if dominated by oaks.

 

Ilex aquifolium is the most familiar member of the genus Ilex, a genus that comprises over 400 species within the family Aquifoliaceae[v]. The holly is a dioecious[vi], broadleaf evergreen native to the understorey of temperate European forests. Holly is recognised for its glossy, spiny leaves and bright red berries, which have inspired wide-ranging cultural and medicinal uses. While prized ornamentally and ecologically in Europe, holly is increasingly problematic in non-native ecosystems, such as in the Pacific Northwest of North America[vii]. I have not had great success when transplanting hollies, but when left to their own devices, the plant can go crazy.


Botanical Description

Ilex aquifolium typically reaches 10–25m in height and displays what is known as significant phenotypic plasticity. This is the ability of an organism to change in response to stimuli or inputs from the environment[viii]. It grows slowly, at around 10-15cm each year[ix] and can live for up to 300 years[x]. Juvenile and lower canopy leaves tend to be spiny, whereas upper canopy leaves often lack spines[xi]. The species produces small, white, four-lobed flowers on separate male and female plants in the early summer. Pollination is insect-mediated, primarily by bees[xii]. Female trees yield fleshy red berries (drupes), each containing 3–4 seeds, which ripen in autumn and persist through winter, providing key food for frugivorous birds such as the blackbird (Turdus merula) and robin (Erithacus rubecula)[xiii].

 

A holly leaf - spiny to keep animals away
A holly leaf - spiny to keep animals away

As an evergreen, the holly remains green and functional throughout the year. However, evergreen trees also lose leaves, but each tree loses its leaves gradually, not all at once. For example, most tropical rainforest plants are evergreens, replacing their leaves gradually throughout the year as the leaves age and fall. Meanwhile, deciduous trees shed their leaves, usually as an adaptation to a cold or dry/wet season[xiv]. Generally, deciduous trees exhibit higher rates of photosynthesis per unit of leaf mass compared with evergreens, especially during favourable conditions. However, over a year, conifers generally photosynthesise more than deciduous trees because they can photosynthesise year-round, while deciduous trees cannot photosynthesise once they lose their leaves. 

 

Ecological Role

1.     Woodland Understorey Dynamics

In native woodlands, holly contributes significantly to vertical stratification and biodiversity. Vertical stratification is a widespread phenomenon in plant and animal communities in woodlands and is a key factor for structuring species richness and biodiversity, particularly in tropical forests[xv].  Holly often dominates the shrub layer of ancient woodlands, particularly in Atlantic oak forests of the UK and the Iberian Peninsula[xvi]. Its dense foliage offers year-round refuge for small mammals and birds. 

Holly can dominate the woodland understorey
Holly can dominate the woodland understorey

2.     Avian and Invertebrate Interactions

Holly berries are a winter food source for birds, although they are toxic to humans and some mammals thanks to the presence of saponins[xvii]. Holly flowers support a diverse invertebrate community, including pollinators and leaf-mining moths such as Phyllonorycter ilicifoliella[xviii], and leaf-mining flies such as Phytomyza ilicis[xix].

 

3.     Soil and Hydrology

The holly’s root system contributes to soil stabilisation and water retention. Its slow decomposition rate and evergreen nature influence nutrient cycling in mixed deciduous forests[xx].

 

Chemical Composition and Pharmacological Potential

Holly leaves contain a range of secondary metabolites with potential pharmacological applications. These include:

 

  • Triterpenoids: Ursolic acid and oleanolic acid exhibit antiviral and anti-inflammatory properties[xxi].

  • Phenolic Compounds: Caffeoylquinic acids contribute to antioxidant activity[xxii].

  • Saponins: Antimicrobial activity has been documented in leaf extracts[xxiii].

 

It appears that there are significant antiviral effects of holly leaf extracts against HSV-1 and adenovirus in vitro, which suggests future medicinal applications[xxiv].

 

Ethnobotanical and Ethnomedical Significance

1.     Cultural and Religious Uses

In Celtic and Roman traditions, holly symbolised eternal life and protection. It was central to winter solstice festivals such as Saturnalia (17-23 December). In Christianity, the red berries and sharp leaves symbolised Christ's blood and the crown of thorns[xxv]. Holly remains a common Christmas decoration across Europe and North America.

Holly's red berries and sharp leaves were said to symbolise Christ's blood and crown of thorns
Holly's red berries and sharp leaves were said to symbolise Christ's blood and crown of thorns

Holly’s symbolic and mythological features have not been comprehensively documented in the academic literature, but it is clear that it has a detailed mythology associated with it. These include:

 

·      Holly in Prehistoric and Celtic Traditions

The veneration of holly in prehistoric Europe is closely associated with the Celtic calendar and Druidic ritual practices. Druids believed holly to be sacred, associating it with protection against evil spirits and misfortune[xxvi]. In the Celtic tree calendar, holly corresponded to the seventh lunar month (typically July), representing masculine energy and the waning year. Evidence from Iron Age archaeological sites suggests that holly branches were used decoratively in winter solstice ceremonies[xxvii]. The persistence of holly leaves in the darkest months served as a powerful symbol of life overcoming death. Additionally, the wood's density and strength made it suitable for constructing weapons and charms, reinforcing its association with strength and protection.

 

·      The Oak King and Holly King Duality

A significant element of Celtic-inspired neopagan mythology was the seasonal battle between the Oak King and the Holly King, which reflected the solar cycle. The Holly King reigned from the summer solstice (21 June) to the winter solstice (21 December), symbolising the descent into darkness, while the Oak King governed the brighter half of the year. Although not part of early Celtic mythology per se, this belief became important in Wiccan[xxviii] and Neopagan[xxix] traditions[xxx]. The Holly King has been depicted as a wintery figure, crowned with holly leaves and armed with a holly club[xxxi], representing introspection and an internal journey.

 

·      Roman Mythology and Saturnalia

The Romans associated holly with Saturn, the god of agriculture and time. During Saturnalia holly wreaths were exchanged as symbols of good fortune and protection[xxxii]. Holly was planted near dwellings to honour Saturn and to repel lightning and malevolent forces.

 

Ovid’s Fasti references the use of evergreen boughs during Saturnalia[xxxiii], although holly is not mentioned by name. However, archaeological and literary evidence confirms its symbolic presence in Roman midwinter festivals[xxxiv].

 

·      Norse Mythology and Germanic Traditions

In northern Europe, holly was linked with the thunder god Thor and thought to protect against lightning strikes. The etymology of "holly" in Old English (holegn) shares roots with words for "holy," and suggests sacred connotations[xxxv]. Germanic tribes used holly in Yule celebrations (21 December – 1 January), a midwinter festival marking the rebirth of the sun. As part of Yule, holly and ivy were brought indoors to symbolise the continuation of life[xxxvi].

 

·      Christian Symbolism

As Christianity spread through Europe, it absorbed many pagan symbols and rituals, including holly. Holly was said to symbolise Christ’s crown of thorns, while the red drupes represented drops of his blood shed during the Passion. The Passion of Christ is the story of Jesus’ arrest, trial, and suffering. It ends with his execution by crucifixion. There is also the English carol "The Holly and the Ivy", first recorded in the 18th century, that places holly and ivy together to express Christian themes of sacrifice and rebirth[xxxvii]. Churches across medieval Europe incorporated holly into their seasonal decorations, and it was often forbidden to remove holly trees from churchyards because of their perceived sanctity[xxxviii].

 

·      Folklore, Superstition, and Rural Beliefs

Holly played a role in many European folk traditions. In English folklore, it was considered bad luck to fell a holly tree. It was said that the tree could ward off witches and evil spirits[xxxix]. Holly branches were commonly hung over doors and windows to protect households during Yule. In Scottish and Irish traditions, holly was associated with the "fairy folk." It was believed that disturbing a holly tree could provoke supernatural retribution[xl]. Holly was also a traditional component of solstice fire rituals, used to banish the old year and invite in the new one.

·      The Holly Tree in Literature and Art

The holly tree has appeared as a literary theme in British Romantic poetry. For example, Robert Southey’s 1809 poem[xli] “The Holly Tree used the tree as a symbol of constancy and endurance. Holly also appeared in Charles Dickens’ 1843 “A Christmas Carol” [xlii] and was related to redemption and spiritual renewal. Holly also featured prominently in medieval and Renaissance art, appearing in nativity scenes and illuminated manuscripts to symbolise divine protection.

First edition of Dickens' "A Christmas Carol" (courtesy Wikipedia)
First edition of Dickens' "A Christmas Carol" (courtesy Wikipedia)

·      Contemporary Paganism and Revivalist Traditions

Modern neopagan and Druidic traditions have revived holly’s mythological associations. Contemporary Wiccan solstice rituals often include holly to represent the waning year or the Holly King[xliii]. Organisations such as the Order of Bards, Ovates, and Druids (OBOD)[xliv] encourage the planting and preservation of holly as part of their ecological and spiritual practice. Holly is a guardian of thresholds - for example, between light and dark, or life and death. The tree remains potent in modern mythology[xlv].

 

2.     Traditional Medicine

Traditional remedies used holly leaves as diuretics and in treatments for joint pain and fevers. Decoctions were made from dried leaves, although excessive use was avoided because of the potential toxicity of berries[xlvi]. Decoction is the action or process of extracting the essence of something.

 

·      Diuretic and Fever Remedies

The most traditional use of holly leaves is as a diuretic and fever reducer (febrifuge). In British and European folk medicine, infusions made from dried holly leaves were consumed to promote urination (diuresis) and to reduce fevers associated with common infections and inflammatory conditions. The 17th-century English herbalist, Nicholas Culpeper, noted that the leaves "provoke urine" and were “good for the stone and gravel in the kidneys"[xlvii].

 

·      Anti-Rheumatic and Anti-Inflammatory Use

Holly leaves were also used externally in poultices to treat joint pain and inflammation, particularly in cases suspected to be rheumatism or gout. Traditional texts described applications where crushed holly leaves were mixed with vinegar and applied to inflamed joints. This may be related to current knowledge that anti-inflammatory triterpenoids can be found in Ilex aquifolium[xlviii].

 

·      Treatment of Digestive and Respiratory Ailments

Less commonly, holly was employed in the treatment of respiratory complaints. In some regions of Spain and France, decoctions were taken to alleviate coughs and congestion[xlix]. Holly was also used to stimulate digestion and appetite after illness. These applications reflect the general use of bitter plant compounds to support gastrointestinal function in traditional European medicine.

 

·      Use of Berries in Folk Medicine

Despite their known toxicity, holly berries have occasionally appeared in folk remedies. In small doses, they were reportedly used as a purgative (laxative) or emetic. However, thanks to saponins, the berries are potentially poisonous, especially to children and pets, so such use is now strongly discouraged [l].

 

Invasiveness and Conservation Concerns

Holly has naturalised in New Zealand, Australia, and the western United States. It sometimes invades native coniferous forests and can displace understorey flora[li]. It does the same in the UK. If it is thought that holly is creating damage, it can be manually removed. However, this is labour-intensive and often followed by resprouting. Chemical treatments with, dare I say it, glyphosate (I hate the stuff) are effective but can be worrying in sensitive habitats[lii].


Holly has naturalised in New Zealand, Australia, and the western USA
Holly has naturalised in New Zealand, Australia, and the western USA

Genetics

There is a role for epigenetics in the leaf variability of Ilex aquifolium. Epigenetics is the study of changes in organisms caused by modification of gene expression rather than alteration of the genetic code itself. It has been found that DNA methylation differences correspond with leaf spine development, allowing holly to adapt to herbivore pressure across its lifespan[liii]. This is epigenetics in action. There are plenty of sheep and deer in the Lake District, and they appear to eat just about anything. Consequently, a holly needs to gather as much protection as it can, so epigenetics are important.

 

Conservation and Legal Status

In the UK, holly is a protected species under the Wildlife and Countryside Act (1981)[liv]. It is listed as a priority species in several ancient woodland conservation plans. Meanwhile, in certain parts of the USA, it is designated as a noxious weed[lv].

 

Holly and the Environment

Ilex aquifolium has been an integral part of European flora for centuries. Its ability to thrive in diverse habitats, from woodlands to urban areas, underscores its adaptability. It is also resilient to climate change. As an evergreen with drought-tolerant features, it may actually benefit from warming temperatures. Its range is expanding northwards in Europe, and its photosynthetic activity in winter gives it a competitive edge over deciduous species[lvi].

 

This adaptability also raises ecological concerns, especially when holly exhibits invasive tendencies outside its native range. The tree can thus both benefit and disadvantage the environment, as follows:

 

Environmental Benefits

  1. Biodiversity Support

Holly trees contribute to biodiversity by providing habitat and food sources for various species:

  • Avian Species: The berries of Ilex aquifolium are a vital food source for birds, especially during winter when other resources are scarce. This facilitates rapid seed spread into remote areas.

  • Invertebrates: The dense foliage of holly offers shelter for insects, including pollinators such as bees and butterflies, which use holly flowers for nectar. ​

Holly everywhere, yet not a drupe to find - mind you, this photograph was taken in March
Holly everywhere, yet not a drupe to find - mind you, this photograph was taken in March

 

  1. Erosion Control

The extensive root system of holly trees stabilises soil and reduces erosion, particularly on slopes and riverbanks. This characteristic is beneficial in maintaining soil integrity and preventing landslides.

 

  1. Carbon Sequestration

As an evergreen species, holly photosynthesises throughout the year, sequestering carbon dioxide and contributing to carbon storage. While individual trees may sequester modest amounts, collectively they play a role in mitigating climate change.

 

  1. Aesthetic and Cultural Value

Beyond ecological functions, holly trees enhance landscape aesthetics, especially during winter months, and hold cultural significance in various traditions, promoting conservation awareness.

 

Environmental Disadvantages

1.     Invasive Potential

While native to parts of Europe, Ilex aquifolium has shown invasive behaviour in several parts of the world. It does this in a number of ways:

 

  • Ecosystem Disruption: In non-native areas, holly can form dense thickets, outcompeting indigenous vegetation, reducing light availability and altering forest understorey composition. This leads to reduced native biodiversity and changes in habitat structure. 


  • Fire Hazard: Holly foliage is highly flammable[lvii] and increases the risk of forest fires. ​


  1. Allelopathic Effects

Some studies suggest that holly may release compounds into the soil that inhibit the growth of surrounding vegetation[lviii]. Allelopathy is a biological phenomenon where one organism releases chemicals (allelochemicals) that influence the growth, survival, and reproduction of other organisms.

A hollinus
A hollinus

  1. Human and Animal Health Concerns


  • Toxicity: Holly berries contain compounds such as theobromine and saponins[lix], which are toxic to humans and pets if eaten. 

  • Physical Harm: The spiny leaves can cause physical injury to humans and animals, deterring movement through dense holly thickets.​

 

Consequently, next time I encounter a holly, I must see it through different eyes. It may be lonely, positioned high on a distant hillside, or among its kind in the centre of dense woodland. Yet it has been, and remains, a valuable plant. Beneficial environmentally, although sometimes a problem, functional medicinally, but also an occasional concern, holly definitely forms part of mythology and history. It may stand quietly, immobile, trying to pretend it is somewhere different. But it is not. Holly, Ilex aquifolium, plays a significant part in our environment, life, and well-being. Never ignore a holly.

 

***

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References


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