Aquifoliaceae Bercht. & J.Presl

First published in Prir. Rostlin Aneb. Rostl. 2(109*): [438], 440. 1825 (1825)nom. cons.
This family is accepted

Descriptions

Aquifoliaceae, E. J. Mendes. Flora Zambesiaca 2:2. 1966

Morphology General Habit
Trees or shrubs, mostly evergreen
Morphology Leaves
Leaves alternate, simple; stipules absent or very small
Morphology Reproductive morphology Flowers
Flowers frequently dioecious, actinomorphic, cymose, fasciculate, subumbellate or rarely solitary
Morphology Reproductive morphology Flowers Calyx
Calyx-lobes imbricate
Morphology Reproductive morphology Flowers Corolla
Petals free or connate at the base, imbricate or valvate
Morphology Reproductive morphology Flowers Androecium
Stamens free, alternate with the corolla-lobes; anthers 2-thecous, opening lengthwise
Morphology Reproductive morphology Flowers Nectaries
Disk absent
Morphology Reproductive morphology Flowers Gynoecium
Ovary superior, 3- or more-locular; style terminal or stigma sessile; ovules 1–2 in each loculus, pendulous from the apex
Morphology Reproductive morphology Fruits
Fruit drupaceous
Morphology Reproductive morphology Seeds
Seeds with copious fleshy endosperm and small straight embryo
[FZ]

Aquifoliaceae, B. Verdcourt. Flora of Tropical East Africa. 1968

Morphology General Habit
Trees or shrubs, mostly evergreen, usually practically glabrous
Morphology Leaves
Leaves alternate or rarely opposite, simple; stipules minute or absent
Morphology Reproductive morphology Flowers
Flowers regular, hermaphrodite or sexes separate, in cymes or fascicles, rarely solitary Male flowers with 4–5(–9) stamens, free or inserted at the base of the corolla-tube, alternating with the petals, hypogynous; anthers dithecous, opening by longitudinal slits; rudimentary ovary 1-locular Female flowers: ovary superior, (2–)3(–22)-locular; ovules 1–2 in each locule, pendulous; style terminal or absent; stigma capitate or discoid; staminodes present but anthers sessile or on shorter filaments
Morphology Reproductive morphology Flowers Calyx
Calyx-lobes 4–6(–9), imbricate
Morphology Reproductive morphology Flowers Corolla
Corolla rotate; petals 4–5(–9), free or joined at the base, imbricate or valvate; disk absent
sex Male
Male flowers with 4–5(–9) stamens, free or inserted at the base of the corolla-tube, alternating with the petals, hypogynous; anthers dithecous, opening by longitudinal slits; rudimentary ovary 1-locular
sex Female
Female flowers: ovary superior, (2–)3(–22)-locular; ovules 1–2 in each locule, pendulous; style terminal or absent; stigma capitate or discoid; staminodes present but anthers sessile or on shorter filaments
Morphology Reproductive morphology Fruits
Fruit a drupe with (1–)2–6(–many) 1(–2)-seeded pyrenes
Morphology Reproductive morphology Seeds
Seeds with copious endosperm
[FTEA]

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit
Trees or shrubs, usually evergreen, sometimes deciduous, rarely prostrate subshrubs (Ilex prostrata Groppo)
Morphology Leaves Stipules
Stipules present, small and often deciduous
Morphology Leaves
Leaves alternate, rarely (sub)opposite, simple, usually glabrous, when pubescent, then the trichomes simple, lamina in some species black-doted, margins entire, crenate, serrate or spinescent
Morphology Reproductive morphology Inflorescences
Inflorescences axillary, derived from thyrses: cymes, thyrsoids, fascicles or apparently solitary flowers
Morphology Reproductive morphology Flowers
Flowers 4-6(-23)- merous, actinomorphic, unisexual by abortion (plants dioecious), sepals of valvate aestivation, little joined at base; petals usually white or cream, base connate to the base; stamens as many as the petals, alternate, reduced to staminodes in the female flowers; gynoecium syncarpic, with 4-6(-23) carpels, ovary superior, locule as many as the carpels, usually with one ovule each, placentation axillary, the ovule inserted at the top of the axis, style very short to absent, stigma flat or discoid; gynoecium reduced to a pistillode in the male flowers
Morphology Reproductive morphology Fruits
Fruit a drupe, with 1-6 pyrenes, with one seed each, epicarp red, purple or virtually black, mesocarp fleshy
Morphology Reproductive morphology Seeds
Seeds with oily, proteinaceous, abundant endosperm .
Note
Number of genera: One genus, Ilex, worldwide, with more than 500 species, ca. 300 of them in Tropical America. Nemopanthus Raf. (one species in North America) was recently included under the synonymy of Ilex (Powell et al. 2000). "Several species of Ilex (e.g. I. aquifolium L., I. cornuta Lindl. & Paxton, I. crenata Thunb., and I. opaca [Soland.]) are cultivated as ornamentals, mainly because their decorative fruits and/or foliage. However, their use in the Neotropics as ornamentals is occasional. The species with most economic importance in the Neotropics is by far Ilex paraguariensis A.St.-Hil., native in Argentina, Brazil, Paraguay, and Uruguay. From the leaves of this species is made a tea, the ""mate"", ""erva-mate"" or ""yerba-mate"", largely consumed in South Brazil, Argentina, Paraguay and Uruguay. Mate is a part of the popular culture in those places, being characteristically served hot in a gourd (called ""cuia"" in Brazil) and drank with a straw (the ""bomba""). It can be also be served as a cold beverage (""tererê""), usually in a decorative cow horn, especially in Paraguay and Brazil. Other species of llex, such as I. dumosa Reissek or I. theezans Mart. were sometimes used in the past as substitutes of the true mate. Other species, as I. tarapotina Loes. (Colombia and Peru) and I. vomitoria Aiton(Mexico and North America) are used as teas, mainly because their emetic or stimulating effects. The wood of some species (I. affinis Gardn., I. brevicuspis Reissek, I. cerasifolia Reissek, I. dumosa,  and I. theezans) is locally used in Brazil in works of carpentry, and as fuel."
Diagnostic
Trees or shrubs with simple usually alternate leaves. Flowers usually white or cream, small, with joined petals at base. Ovary syncarpic with short or absent style. Fruit a red, purple or virtually black drupe with 4-6 pyrenes. Although the plants are dioecious, the presence of staminodes in the pistillate flowers and of a pistillodium in the staminate flowers may misinterpret the flowers as bisexual. Key differences from similar families: Aquifoliaceae is frequently confused with Celastraceae (Maytenus Molina spp.) and Symplocaceae (Symplocos Jacq.). From Maytenus it is readily distinguished by the lack of a disc in the flowers (conspicuous in Maytenus), and the fruit (drupe vs. capsule in Maytenus). Leaves in species of Ilex are not pruinose (covered with a wax layer) like in many species of Maytenus. Additionally, the top of the branches in Maytenus is, in many species, 4-angled or striate, or even winged. In Ilex the branches are usually rounded in transverse section. Symplocos species often bear flowers with 5-100 stamens (in 1-4 whorls), with a nectariferous disc surrounding the style base (absent in Ilex). The style is clearly distinct from the ovary (short or absent in Ilex), this (sub) inferior (superior in Ilex). Leaves in Symplocos usually become light-green when dry, different from Ilex (grayish or dark-colored). Additionally, many species of Symplocos have glandular-denticulate leaves. Distinguishing characters (always present): Leaves simple. Plants dioecious; female flowers with staminodes, male flowers with pistillode. Flowers isostemonous, stamens alternate to the petals; petals joined at base. Ovarysuperior. Short or absent style. Fruit a drupe, usually with 4-6 pyrenes. Leaves alternate, seldom (sub)opposite. Stipules small, often deciduous. Flowers small, actinomorphic, usually white or cream. Drupe red, purple or virtually black.
Distribution
Ilex (Native) In the Neotropics, Ilex L. can be found from sea level up to ca. 4,000 metres (Andes).
[NTK]

Aquifoliaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Morphology General Habit
Trees or shrubs, mostly evergreen
Morphology Leaves
Leaves alternate, simple; stipules absent, or rarely present
Morphology Reproductive morphology Flowers
Flowers actinomorphic, hermaphrodite or unisexual, fasciculate or subumbellate or rarely solitary
Morphology Reproductive morphology Flowers Calyx
Calyx-lobes imbricate
Morphology Reproductive morphology Flowers Corolla
Petals 4 or 5, free or connate at the base, hypogynous, imbricate
Morphology Reproductive morphology Flowers Androecium
Stamens hypogynous, 4 or 5, alternate with the petals, rarely more numerous, free; anthers 2-celled, opening lengthwise
Morphology Reproductive morphology Flowers Nectaries
Disk absent
Morphology Reproductive morphology Flowers Gynoecium
Ovary superior, 3- or more-celled; style terminal or absent; ovules 1–2 in each cell, pendulous from the apex
Morphology Reproductive morphology Fruits
Fruit drupaceous, of 3 or more 1-seeded pyrenes; seed with copious fleshy endosperm and small straight embryo
[FWTA]

Timothy M. A. Utteridge and Laura V. S. Jennings (2022). Trees of New Guinea. Kew Publishing. Royal Botanic Gardens, Kew

Ecology
Monotypic family with c. 600 species, throughout the tropics, subtropics and temperate regions; at least 13 species in New Guinea.
[TONG]

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition
Characters of similar families: Berberidaceae: stipules usually absent, flowers bisexual, petals with nectaries, fruits berries. Celastraceae: flowers bisexual, fruits berries or woody capsules. Fagaceae (Quercus ilex): flowers unisexual, monoecious, fruit a single-seeded nut in cupule. Lauraceae: often aromatic, stipules absent, anthers with flaps, fruit a berry in cupule. Oleaceae (Osmanthus spp.): opposite leaves, flowers usually bisexual, sepals usually fused into a tube. Symplocaceae (Symplocos spp.): stipules absent, stamens numerous, fruit often blueish.
Morphology General Habit
Trees and shrubs, evergreen or deciduous
Morphology Leaves Stipules
Stipules small, persistent or caducous leaving a scar
Morphology Leaves
Leaves simple, alternate, rarely opposite or subopposite; leaf blade usually coriaceous, sometimes chartaceous or membranous, margin entire to spinose toothed
Morphology Reproductive morphology Inflorescences
Inflorescence axillary, solitary, fasciculate or rarely in cymes
Morphology Reproductive morphology Flowers
Flowers usually dioecious, actinomorphic, small, isomerous; calyx 4–5(8)-lobed, persistent, coriaceous, green; petals 4–5(8), imbricate, mostly fused at base or up to half of length, often white, cream, green, yellow, pink, or red; stamens or staminodes alternating with petals, epipetalous
sex Male
Male flowers: anthers oblong-ovoid, introrse, longitudinally dehiscent; ovulode, subglobose or pulvinate, rostrate
sex Female
Female flowers: staminodes sagittate or cordate; ovary superior, ovoid, 4–8(–10)-loculed, rarely pubescent; style very short to absent; stigma capitate, discoid, or columnar
Morphology Reproductive morphology Fruits
Fruit a drupe, red, brown, black, green or yellow, globose; stigma persistent, conspicuous; exocarp chartaceous or membranous; mesocarp fleshy
Morphology Reproductive morphology Seeds
Seeds pyrenes (1–)4–6(–23).
Distribution
A family of 1 genus and ca. 500–600 species; worldwide with centres of diversity in South America and Southeast Asia.
Note
Trees or shrubs, dioecious. Stipules small. Leaves simple, usually alternate and leathery, margins entire to spinose. Flowers unisexual; actinomorphic, isomerous, inconspicuous, unisexual, ovary superior. Fruit a drupe, usually globose.
Description Author
Sara Edwards
[KTEMP-FIH]

Uses

Use
Ilex paraguayensis is used to make maté tea, which is commonly drunk in South America. Other species of Ilex are used as ornamental plants. The wood is used in carving and furniture-making as it is unusually pale.
[KTEMP-FIH]

Sources

  • Flora Zambesiaca

    • Flora Zambesiaca
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Flora of Tropical East Africa

    • Flora of Tropical East Africa
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Flora of West Tropical Africa

    • Flora of West Tropical Africa
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Herbarium Catalogue Specimens

    • 'The Herbarium Catalogue, Royal Botanic Gardens, Kew. Published on the Internet http://www.kew.org/herbcat [accessed on Day Month Year]'. Please enter the date on which you consulted the system.
  • Kew Names and Taxonomic Backbone

    • The International Plant Names Index and World Checklist of Vascular Plants 2025. Published on the Internet at http://www.ipni.org and https://powo.science.kew.org/
    • © Copyright 2023 International Plant Names Index and World Checklist of Vascular Plants. http://creativecommons.org/licenses/by/3.0
  • Neotropikey

    • Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • Trees of New Guinea

    • Trees of New Guinea
    • http://creativecommons.org/licenses/by-nc-sa/3.0
  • The Kew Temperate Plant Families Identification Handbook

    • The Kew Temperate Plant Families Identification Handbook
    • http://creativecommons.org/licenses/by-nc-sa/3.0