Order Santalales / Sandalwood & Mistletoe Flowers

    Order Santalales

    Santalales is one of the most prominent groups of parasitic flowering plants, with species distributed worldwide. Santalales involve plants that form haustoriumWhat is haustorium?A specialized root of parasitic vascular plants capable of penetrating and absorbing food and other materials from host tissues. to penetrate the host’s barks and roots, plants that undergo photosynthesizing activities in the stem, and members with usually simple and exstipulateWhat is exstipulate?Without stipules; Stipule is a small structure of appendage found at the base of some leaf petioles. leaves.

    The Santalales flowers are bisexual or unisexual and are generally pollinated by nectar-foraging birds, moths, bees, wasps, and flies. Sandalwood and mistletoes are just two of the known plants under Santalales.

    Santalales Families

    Many studies present different quantities of families under Santalales. But due to the differences in sampling results, the relationship between the families and even which families should be placed in this order is still not fully determined[1]. For example, the APG IV system (2016) accepted seven families belonging to Santalales. This value is also used in the World Flora Online[2].

    • Balanophoraceae (Snake-mushroom family)
    • Loranthaceae (Showy mistletoe family).
    • Misodendraceae (Feathery-mistletoe family)
    • Olacaceae (Tallow-wood family)
    • Opiliaceae (Bally-coma family
    • Santalaceae(Sandalwood family)
    • Schoepfiaceae (Arana family)

    Santalales Distribution

    Santalales Distribution

    The species of Santalales are distributed worldwide. However, its heavy concentration is found in tropical and subtropical regions.

    • The species Santalaceae[3] (40 genera and 990 species[4] )are found worldwide. Species are found in the dry or temperate regions or the humid tropical biomes.
    • Loranthaceae (89 genera and 886 species) are scattered worldwide. The Loranths are incredibly diverse in Africa and Australia’s seasonally dry habitats. A few species of this family also exist in temperate Europe, Asia, Australia, New Zealand, and South America.
    • The Olacaceae[5] (27 genera and 149 species) has a pantropical distribution. However, members are also found in subtropical North and South America. The plants under this family can survive under low nutrients and water by attaching to their hosts during dry periods.
    • The Opiliaceae[6] (10 genera and 33 species) members are also pantropical, extending their distribution to temperate Australasia. They are mainly distributed in the Old World. Only one genus, Agonondra, is found in the Neotropics.
    • Balanophoraceae (16 genera and 45 species) have a tropical to subtropical distribution. They are also found in South Africa, the Himalayas, China, Japan, and New Zealand.
    • The members of Misodendraceaea (1 genus and 8 species) are widespread. This family is more diverse in the Southern hemisphere, chiefly in Australia. However, diversity in flower morphology is also found in South America.

    Santalales Characteristics

    Santalales Characteristics

    Below are some of the characteristics shared by the members of Santalales:

    • Plant type: Santalales members are trees, lianas, shrubs, herbs, and terrestrial and evergreen vines. They are characterized as parasitic. Most of them are hemiparasitesWhat is hemiparasites?A parasitic plant containing chlorophyll and therefore partly self-sustaining as in mistletoe.. They can undergo photosynthesis and produce sugars, but they get water and minerals by attaching them to the roots or stems of other plants. Somespecies are holoparasitesWhat is holoparasites?A plant lacking chlorophyll and entirely dependent on its host for nourishment. They are subdivided into stem or root parasites depending on which part of the host they attack. and depend on a significant part of their nourishment from their hosts.
    • Roots: In Santalaceae, the roots are present or absent. The mistletoe after germination, uses the modified root, haustorium, to penetrate the bark or the root of the host.
    • Stem: Surface hairs are found on the stems of some species. Some species transfer the photosynthesizing function to the stem.
    • Leaves: Almost all species have simple leaves; exstipulate. There are surface hairs on a few species’ leaves.
    • Flowers and Inflorescences: Variation in the inflorescence and flower morphology in this order is extreme. (Please see the flowers and reproduction and family differences section of this page).
    • Sepals and Petals: Some flowers have both sepals and petals. Others have either or none at all.
    • Stamens and Carpels: The stamens are found opposite the petal’s lobes and are typically equal to the quantity of the petals. The carpels are usually 2-5.
    • Ovary and Fruits: The ovaries are superior, partly inferior, or inferior (mostly). The species have remarkable ovule, embryo sac, and embryo development. The Loranthaceae do not have integuments in the ovules. Other families have pendulous ovules in the free central placenta. The fruit is a berry, drupe, nut, or achene.
    • Seeds: The seeds of the members mostly lack testae (seed coats).

    Santalales Flowers and Reproduction

    Santalales Flowers and Reproduction

    The Santalaceae flower size varies considerably. Most species are small and inconspicuous. Although some members (Santalum and Quinchamalium) have showy flowers of moderate size.

    Generally, the Santalales species[9] are pollinated by nectar-foraging birds, moths, bees, wasps, and flies. The birds, wind, and explosion are the means to disperse the fruit or seed.

    Santalales Family differences

    The Sandalwood[10] is an order of parasitic flowering plants with exceptionally diverse growth habits. There are holoparasites and depend mostly on their hosts. However, most species have at least some chlorophylls and can still undergo photosynthesis.

    The following are the differences of the principal families (families with the most number of genera):




    • The members are fleshy, herbaceous parasites that grow on the roots or rhizomes of their hosts. The leaves are scale-like, alternate, and spirally arranged.
    • The plants under this family lack chlorophyll and stomata. The species form an underground tuber where the rhizomes grow, forming haustoria that attach to the host’s roots.
    • The flowers are tiny, unisexual, and actinomorphic. The inflorescences are in terminal racemes that are reduced to spadices. These inflorescences consist of unisexual flowers or both male and female flowers.
    • The female flowers are smaller than the male flowers. In male flowers, the petals are lacking. The free or fused sepals can be 3 or 4, rarely 8. Some species also lack sepals.
    • The female flowers are crowded. As a result, both the sepals and petals are absent. If present, they are reduced to 2 free or fused tepals.
    • The stamens are as many as the sepals. If the sepals are absent, the stamens are 1-2.
    • The ovary is inferior with 2-3 fused carpels.
    • The fruit is an achene.


    • Members of this family are terrestrial and evergreen vines, shrubs, and trees. Mostly, they are green and autotrophic. However, some members are parasitic that attach to their hosts through their haustorical connections.
    • The leaves are simple, alternate, distichousWhat is distichous?Two-ranked, with leaves or flowers in two opposite rows in the same plane., or arranged spirally. The petioles are well developed. The stipules are lacking. Their leaf blade possesses pinnate venation, and the leaf margins are entire or toothed.
    • The flowers are actinomorphic and bisexual or partially unisexual. They are in inflorescences of cymes, racemes, fascicles, spikes, or umbels (simple or compound).
    • There are usually 5 fused sepals forming a cup shape. However, this calyx is sometimes absent in some species. The petals are also five, free or slightly fused.
    • The stamens are commonly 4-5, sometimes 8-10; in 1 or 2 whorls.
    • The ovary is superior or half inferior. The carpels are 3-5; fused.
    • The fruits are drupes.

    Santalales Example species

    The species under this order are mostly parasitic. However, these plants still provide beneficial uses. The following are some of the example species of Santalales:

    • Aromatic Sandalwood – The species is economically important in perfumery and furniture making.
    • Fungus root – The plant has medicinal use.
    • Moodjar or Western Australian Christmas tree[11] – The plant has potential for medicine.
    • Large sourplum – The plant is beneficial. It can be a source of food (fruit), oil (from seeds), firewood (from wood), and medicine[12].
    • Tallowwood[13] – This tree is very beneficial. It is a source of food (fruit), oil (from fruit), essential oils (from heartwood and flowers), and firewood or charcoal (from wood). The tree is also grown as ornamental. It is extensively utilized as a medicinal plant. The leaves are also vital as a source of animal feed.
    • Ivory wood tree[14] – The fruits of this plant can be cooked, and the roots are used to make a frothy drink. The wood is of good quality for export. Aside from being an ornamental plant, it is also a source of fuel and charcoal. Additionally, the parts (leaves, bark, and roots) are of great medicinal value.
    • Reed’s False Olive[15] – The plant is used for traditional medicine. The plant is potentially useful[16] for its anthelminthic activity.
    • Northern Sandalwood – The plant is used as traditional medicine.
    • Bitter quandong – The bark of the roots is edible after roasting.
    • Wood rose[17] – This plant is sometimes used as ornamental. It also has medicinal value.

    Cite This Page

    BioExplorer.net. (2024, June 13). Order Santalales / Sandalwood & Mistletoe Flowers. Bio Explorer. https://www.bioexplorer.net/order-santalales/.
    BioExplorer.net. "Order Santalales / Sandalwood & Mistletoe Flowers" Bio Explorer, 13 June 2024, https://www.bioexplorer.net/order-santalales/.
    BioExplorer.net. "Order Santalales / Sandalwood & Mistletoe Flowers" Bio Explorer, June 13 2024. https://www.bioexplorer.net/order-santalales/.
    Key References
    • [1]“Plants of the World: An Illustrated Encyclopedia of Vascular Plants – Maarten J. M. Christenhusz, Michael F. Fay, Mark W. Chase – Google Mga aklat”. Accessed January 02, 2022. Link.
    • [2]“Search”. Accessed January 02, 2022. Link.
    • [3]“A Molecular Phylogeny of Santalaceae (Santalales) on JSTOR”. Accessed January 02, 2022. Link.
    • [4]“Search”. Accessed January 02, 2022. Link.
    • [5]“Plants of the World: An Illustrated Encyclopedia of Vascular Plants – Maarten J. M. Christenhusz, Michael F. Fay, Mark W. Chase – Google Mga aklat”. Accessed January 02, 2022. Link.
    • [6]“(PDF) Phylogeny and a new tribal classification of Opiliaceae (Santalales) based on molecular and morphological evidence: Phylogeny and classification of Opiliaceae”. Accessed January 02, 2022. Link.
    • [7]“Pollination biology and fertilization in Santalum album L. (Santalaceae) – ScienceDirect”. Accessed January 02, 2022. Link.
    • [8]“American Journal of Botany”. Accessed January 02, 2022. Link.
      by SIDONIE BELLOT AND SUSANNE S. RENNER. Accessed January 02, 2022. Link.
    • [10]“Santalales – an overview | ScienceDirect Topics”. Accessed January 02, 2022. Link.
    • [11]“WA Christmas Tree (Nuytsia floribunda)”. Accessed January 02, 2022. Link.
    • [12]“Ximenia caffra Sond. (Ximeniaceae) in sub-Saharan Africa: A synthesis and review of its medicinal potential – ScienceDirect”. Accessed January 02, 2022. Link.
    • [13]“Ximenia americana: Economic Importance, Medicinal Value, and Current Status in Ethiopia”. Accessed January 02, 2022. Link.
    • [14]“Agonandra brasiliensis – Useful Tropical Plants”. Accessed January 02, 2022. Link.
    • [15]“Cansjera rheedei – Rheed’s False Olive”. Accessed January 02, 2022. Link.
    • [16]“Anthelminthic Activity of Cansjera rheedii J. Gmelin (Opiliaceae)” by V.M. Mounnissamy, S. Kavimani, V. Balu and S. Darlin Quine. Accessed January 02, 2022. Link.
    • [17]“Taranaki Flora – Plants – Dactylanthus taylorii”. Accessed January 02, 2022. Link.


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