Order Aquifoliales / Holly & Brazillian Tea Flowers

Aquifoliales is under the core asterid clade with 5 families and more than 536 species.

• Aquifoliaceae (Holly family)
• Cardiopteridaceae (Chilean citronella tree family).
• Helwingiaceae (Japanese helwingia family).
• Phyllonomaceae
• Stemonuraceae

Aquifoliaceae(1 genus and 400-405 species)^[1] comprises species that are mostly distributed worldwide. This involves the Ilex, including Nemopanthus. On the other hand, Cardiopteridaceae (6 genera and 46 species^[2]) members are mostly found in the tropics.

However, there are also members distributed in temperate regions. Phyllonomaceae (1 genus and 5 species)^[3] are found in Asia, northern Indochina, and the Himalayas. Phyllonoma, the sole genus of Phyllonomaceae, is found in South and Central America.

Stemonuraceae (12 genera and about 90 species)^[4] are chiefly distributed in the tropics of the Old World. However, there are also members found in the subtropics.

Members of Aquifoliales share the following characteristics:

• Plant type: Members are usually evergreen trees and shrubs, rarely lianas and climbing herbs.
• Stem: Scalariform perforation plates are usually in the vessel elements of the members. The wood rays are commonly multiseriate and heterocellular.
• Leaves: The leaves of the plants are typically alternate and rarely opposite. They have very small stipules (often caducous). Some species lack the stipules. Members do not possess leaf sheaths. The venation is mostly pinnate and rarely palmate. Resiniferous or laticiferous idioblast are usually found in the mesophylls. Their leaf margins are commonly serrated.
• Flowers and inflorescences: Most of the members are bisexual. A few species are andromonoecious, gynomonoecious, polygamomonoecious, and androdioecious. The flowers are small and actinomorphic. The inflorescences of the flowers are typically axillary, panicle, raceme-like, or fascicle. A few members have solitary flowers. Sometimes, the inflorescence is epiphyllous.
• Sepals and petals: The sepals and petals are usually 4-5; usually connate. The nectaries are typically found at the base of the petals. Some species lack nectaries. The disc is usually absent (poorly developed if present).
• Stamens and carpels: The stamens are commonly 4-5; alternipetalous. There are 2-6 (to 24) carpels; connate. The style is simple and single (or absent). Pistillodium is found in male flowers.
• Ovary and fruit: The ovary is commonly superior and rarely inferior. The placentation of the flowers is typically apical. The fruit is commonly a drupe, rarely a berry or a samara.
• Seeds: The testa of the seed is usually thick; aril absent. Chlorophyll is typically absent in the usually small and straight embryo.

The Holly flowers are small in solitary or branched arrays. They are radially symmetrical (actinomorphic), usually unisexual, and hypogynous. The perianth is dichlamydous.

• Flowers are 4- to 8- 8-parted and possess small sepals surrounding the petals. The sepals and the petals are 4 (-8, 0) and fused at the base. The calyxWhat is calyx?A collective term for all the sepals of a flower; the lowermost whorl of floral orgrans (Plural form is calyces). is synsepalous, and the corollaWhat is corolla?A collective term referring to the petals of a flower. is usually sympetalous. Usually, 4 (-8) stamens grow from the petals’ fused base.
• The ovary is superior, and the gynoecium is syncarpousWhat is syncarpous?Having united carpels. Contrast apocarpous.. The fruit of the Aquifoliales is small and fleshy. The miniature Holly flowers attract bees, butterflies, flies, and other pollinators.
• Cardiopteridaceae flowers are usually small (a few genera have larger flowers) in white, green, yellow, or purple. They are actinomorphic and hypogynous. Most members are bisexual.
• Some species are functionally dioeciousWhat is dioecious?Pertaining to plants, individuals of which bear either staminate or pistillate flowers, but not both. (Gonocaryum) and polygamous (Citronella and Cardiopteris). The species are in the inflorescence of axillary or terminal, cymose, or raceme. Most species have bracts.
• The sepals and petals are usually 4 or 5; usually connate. The petals of Citronella species are not or slightly connate. Flowers have 4 or 5 haplostemonous, antesepalous, alternipetalous stamens. There are 2 to 3 connate carpels in the pistil. The ovary of the species is superior.
• Helwingiaceae flowers are small, actinomorphic, and hypogynous. The male and female flowers are on separate plants (dioecious). Inflorescences of the species are axillary, fasciculate, or epiphyllous (from the upper side of the midrib). The perianth is petaline. Members have 3-5 sepals, valvate to imbricate, and connate. The flowers lack petals. Insect pollination^[5] occurs in this family.

Aquifoliaceae

• Members are trees or shrubs. They are rarely lianas or epiphytes. The leaves are usually alternate, simple, and entire to pinnately lobed. Stipules are minute to absent. The venation is pinnate. Their leaf margin is usually serrate. Other members have dentate or entire. The resiniferous or laticiferous idioblasts are often in the mesophyll.
• Most of the members are dioecious or bisexual. They are rarely polygamomonoecious.
• The flowers are small and actinomorphic in axillary, umbel- or raceme-like, panicle, and fascicle. A few flowers are in solitary axillary.
• The petals and sepals are usually 4 or 5.
• The nectaries are found at the base of the petals.
• The stamens of the flowers are 4, 5, or 8.
• The ovary is superior. There are usually 4 to 6 connate carpels, antepetalous. Pistillodium is present in the male flowers.
• The fruit is a drupe. Aril is absent in seeds. The embryo is minute.

Cardiopteridaceae

• The members are evergreen trees, shrubs, climbing, and twining herbs. The leaves are simple and usually alternate. They are entire or palmately lobed. They lack stipules and leaf sheaths. The venation is typically pinnate, and the leaf margins are primarily entire.
• They are usually bisexual. Some species are andromonoecious, gynomonoecious, and polygamomonoecious. Other species are dioecious, androdioecious, or gynodioecious.
• The flowers are small, actinomorphic, and hypogynous. The flowers are axillary or terminal, cymose or raceme. Members often lack bracts.
• There are usually four or five; connate.
• Most members have 4-5 stamens; antesepalous, alternipetalous.
• The pistil has 2-3 connate carpels. The ovary is superior. Reduced pistillodium is sometimes present in male flowers.
• The fruit is a drupe or two-winged samara. Aril is absent in seeds. The embryo is usually tiny.

Helwingiaceae

• The members are evergreen shrubs or small trees. The leaves are simple, entire, and alternate. The ptyxis is supervolute to curved. The stipules and leaf sheath are absent. The venation of the leaves is pinnate. Their mesophyll lacks calcium oxalate. Their leaf margin is serrated.
• The plants are dioecious.
• The flowers are small, actinomorphic, and hypogynous. The inflorescences are axillary, fasciculate or umbel-like cymose. They are epiphyllous.
• The petals are absent, and the sepals are 3-5; valvate to imbricate, connate.
• There are 3-5 alternipetalous, obhaplostemonous stamens.
• The pistils have 3-5 carpels; connate, anternipetalous. The ovary is inferior. Pistillodium is absent.
• The fruit is a drupe. The seeds have thin testa; aril absent. The embryo is small and straight.

Phyllonomaceae

• The members are evergreen trees or shrubs. The leaves are simple, entire, and usually distichousWhat is distichous?Two-ranked, with leaves or flowers in two opposite rows in the same plane.. They lack leaf sheaths. They have pinnate, brochidodromous venation. Their leaf margin is entire or serrate.
• Members are commonly bisexual.
• The flowers are small, actinomorphic, and epigynous. Flowers are epiphyllous. The bracteoles are absent.
• The sepals are 4-5; free. The petals are 3-5; free.
• There are 4-5 antesepalous, alternipetalous stamens.
• There are commonly 2 connate carpels in the pistil. The ovary is inferior. The pistillodium is absent.
• The fruit is a berry with persistent calyx. The seeds have multilayered testa; aril is absent. The embryo is large and straight.

Stemonuraceae

• Members are evergreen trees or shrubs. The leaves are simple, alternate, and lobate or entire. The ptyxis is conduplicate. They lack leaf sheath and stipules. Members have pinnate, brochidodromous venation. There are sclerenchymatous idioblasts in the mesophyll. Their leaf margin is entire.
• Most members are dioecious. Others are bisexual.
• The flowers are usually small, actinomorphic, and possess an articulated pedicel. They are often axillary in a raceme-like cymose.
• There are usually 4-5 sepals; connate. The petals are commonly 4-5; usually connate, others are free.
• There are typically 4-5 antesepalous, alternipetalous stamens.
• The pistil has 3 (2 or 5) connate carpels. The ovary is superior. Pistillodium is present in the male flowers.
• The fruit is a drupe. The seeds have thin testa; aril is absent. They have small, straight, or curved embryos.

There are more than 536 species under Aquifoliales. The following are the example species under this order.

• Common holly^[6] – This plant is commonly grown in parks and gardens. They are usually used for hedges and in traditional medicine.
• American holly^[7] – This species is an ornamental plant. The wood is used for carpentry, and the leaves can be used to make tea.
• Brazilian tea^[8] – The plant is used for making a beverage. It is also used as a healthy food.
• Chilean citronella tree – The plant has good ornamental value. It also produces oil that repels insects. In addition, the leaves have aromatic and medicinal properties.
• Churnwood^[9] – This grass produces an oil with many benefits. This species is also used as a garden plant.
• Helwingia chinensis – The flowers and leaves of this plant might be edible. The leaves and bark decoction are used for the treatment of skin inflammations.
• Japanese helwingia^[10] – The young flowers and shoots are edible. The leaves can be boiled as a potherb.
• Phyllonoma laticuspis – The leaves of this species have great medicinal value^[11].
• Gomphandra mollis^[12] – The root of this plant is used in traditional medicine.
• Cantleya corniculata^[13] – The fruit is edible. The wood is used for construction and furniture making.