1. Aquifoliaceae Bercht. & J.Presl

    1. This family is accepted.

[FZ]

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

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

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

Habit
Trees or shrubs, mostly evergreen, usually practically glabrous
Leaves
Leaves alternate or rarely opposite, simple; stipules minute or absent
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
Calyx
Calyx-lobes 4–6(–9), imbricate
Corolla
Corolla rotate; petals 4–5(–9), free or joined at the base, imbricate or valvate; disk absent
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
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
Fruits
Fruit a drupe with (1–)2–6(–many) 1(–2)-seeded pyrenes
Seeds
Seeds with copious endosperm
[FWTA]

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

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

Groppo, M. (2009). Neotropical Aquifoliaceae.

Morphology
Description

Trees or shrubs, usually evergreen , sometimes deciduous , rarely prostrate subshrubs (Ilex prostrata Groppo). Stipules present, small and often deciduous . 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. Inflorescences axillary , derived from thyrses: cymes, thyrsoids, fascicles or apparently solitary 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. Fruit a drupe , with 1-6 pyrenes, with one seed each, epicarp red, purple or virtually black, mesocarp fleshy . Seeds with oily, proteinaceous, abundant endosperm .

Distribution
Distribution in the Neotropics
  • In the Neotropics, Ilex L. can be found from sea level up to ca. 4,000 metres (Andes).
Diagnostic
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.
Useful tips for generic identification
  • 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. 
Other important characters
  • Leaves alternate, seldom (sub)opposite.
  • Stipules small, often deciduous.
  • Flowers small, actinomorphic, usually white or cream.
  • Drupe red, purple or virtually black.
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.
General Description
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).
Status
  • Ilex (Native)
General notes
  • 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.
Literature
Important literature

Andrews, S. 1985. A checklist of Aquifoliaceae of Bahia. Rodriguésia 37(63): 34-44.

Baas, P. 1974. The wood anatomical range in Ilex (Aquifoliaceae) and its ecological and phylogenetic significance. Blumea 21: 193-258.

Edwin, G. & Reitz, r. 1967. Aquifoliaceae. In: Reitz, R. (ed) Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí.

Giberti, G.C. 1979. Las especies argentinas del género Ilex (Aquifoliaceae). Darwiniana 22(1-3): 217-240.

Giberti, G.C. 1994. Aquifoliaceae. In: Spichiger, R. & Ramella, L. (eds.) Flora del Paraguay. Conservatoire et Jardin botaniques de la Ville de Genève/Missouri Botanical Garden.

Groppo, M. 2003. Aquifoliaceae. in: Cavalcanti, T.B. & Ramos, A.E. (orgs.) Flora do Distrito Federal, Brasil. v.3: 51-62. Embrapa Recursos Genéticos e Biotecnologia

Groppo, M. & Pirani, J.R. 2002. Aquifoliaceae. In In: Wanderley, M.G.L., Shepherd, G.J., Giulietti, A.M. (Coord.). Flora Fanerogâmica do Estado de São Paulo. vol. 2: 31-37. São Paulo, Fapesp/HUCITEC.

Loesener, T. 1942. Aquifoliaceae. In A. Engler & K. Prantl (Eds.) Die Natürlichen Pflanzenfamilien 2nd ed. 20b: 36-86. Wilhelm Engelamnn, Leipzig.

Loizeau, P.-A. 1994. Les Aquifoliaceae péruviennes. Boissiera 48: 1-306.

Loizeau, P.-A. & Spichiger, R. Aquifoliaceae. In N. Smith, S.A. Mori, A. Henderson, D.W. Stevenson & S.V. Heald (eds.) Flowering Plants of the Neotropics. p. 26-27. Princeton University Press, Princeton and Oxford.

Powell, M., Savolainen, V., Cuénoud, P., Manen J.-F. & Andrews, S. 2000. The mountain holly (Nemophantus mucronatus) revised with molecular data. Kew Bulletin 55: 341-347.

Reissek, S. 1861. Ilicineae. In C.P.F Martius & A.W.Eichler (eds.) Flora brasiliensis. 11(1): 37-80. Monachii. Lipsiae.

Viani, R.A.G. & Vieira, A.O.S. 2007. Flora arbórea do rio Tabagi (Paraná, Brasil): Celastrales sensu Cronquist. Acta Botanica Brasilica 21(2): 457-472.

Images

Aquifoliaceae Bercht. & J.Presl appears in other Kew resources:

First published in Prir. Rostlin Aneb. Rostl. 2(109*): [438], 440. 1825 (1825)

Accepted by

  • APG IV (2016) http://dx.doi.org/10.1111/boj.12385

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

Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2019. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. 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/3.0