Santalaceae R.Br.

Family

Angiosperms > Santalales

Characteristics

Herbs or shrubs, rarely trees, usually root hemiparasites, occasionally aerial hemiparasites (Dendrotrophe and Phacellaria); nodes not articulated, mostly glabrous, less often hairy, hairs simple. Leaves usually alternate (opposite in Buckleya), sometimes scale-like (absent in Phacellaria); stipules absent; petiole often indistinct; leaf blade simple, usually pinnately veined, sometimes palmately 3-9-veined (in Dendrotrophe), margin entire. Inflorescences mostly axillary, occasionally terminal (in Buckleya), cymose, umbellate, paniculate, racemelike, spicate, or clustered, sometimes 1-flowered; bracts scale-like, small, sometimes forming involucre, sometimes ± adnate to pedicel (in Thesium); bracteoles sometimes present, paired. Flowers bisexual or unisexual (plants usually dioecious, rarely monoecious), actinomorphic, 3-6(-8)-merous, very small; perianth lobes 3-6(-8), slightly fleshy. Male flowers: perianth lobes valvate or slightly imbricate in bud, incurved or patent when flowering, sparsely hairy or with tongue-shaped appendage at the insertion of the stamens. Disk epigynous or perigynous, sometimes absent, margin sinuate or lobed, sometimes distinct, glandular or scaly. Stamens as many as and opposite to perianth lobes, usually on the base of lobes; filaments filiform, short; anthers gynobasic or dorsifixed near base, 2-celled, parallel or divaricate, dehiscence usually longitudinal. Female and bisexual flowers: perianth tube usually longer than that of male. Ovary inferior or half-inferior, 1-or 5-12-loculed; ovules 1-3(-5), anatropous or hemitropous, integument absent. Style 1; stigma capitate, small, truncate or lobed. Fruit a drupe or a nut, exocarp usually fleshy, endocarp crustaceous or bony. Seed 1, without a differentiated testa; endosperm copious, usually white and partitioned, fleshy; embryo cylindric, straight, small, smooth, rugose, or many ridged. 2n = 5, 6, 7, 12, 13+.
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Herbs or shrubs, sometimes trees, usually root or stem-hemiparasites, occasionally aerial hemiparasites (Visceae) on stems or branches of the host, bisexual or monoecious or dioecious; glabrous or hairy and the hairs simple or stellate. Haustorium sometimes present (Visceae); haustorial attachment single, epicortical runners absent. Leaves opposite or alternate, rarely whorled, often reduced and scale-like, simple, entire, often soft, sometimes caducous, petiolate; stipules absent. Inflorescence a raceme, spike, panicle, corymb or cluster, often reduced to a single flower, axillary or terminal, usually bracteate, sometimes bracteolate. Flowers actinomorphic, bisexual or unisexual on monoecious or dioecious plants. Perianth usually 1-whorled. Sepals usually absent, or rarely present as a small rim (calyculus) or rarely developed into sepal lobes (e.g. in Buckleya, Osyridocarpos, Thesium libericum, these taxa not in Australia). Petals (2–) 4–5 (–8), rarely more, valvate, free or united into a campanulate or urn-shaped tube. Stamens as many as tepals and opposite them; anthers tetrasporangiate, dithecal or almost so, but often lobed and appearing 4-locular, introrse, dehiscing by longitudinal slits or sometimes by a single apical pore. Gynoecium of 2–5 connate carpels. Ovary inferior, half-inferior or superior, completely unilocular or 1–3-locular basally or 1-locular distally, or the ovary sometimes solid and without differentiated locules. Ovules 1–4 (or ovules sometimes not differentiated); placentation free central. A glandular disc present on top of the ovary, often thickened and lobed at margin, rarely absent; style apical, simple, usually very short; stigma nipple-like or 2-, 3-or 5-lobed. Fruit a nutlet, drupe or pseudodrupe or berries; mesocarp often somewhat fleshy; receptacle often enlarged, fleshy and coloured. Seed 1, without testa; endosperm usually copious.
Herbs, shrubs or trees, hemiparasitic. Leaves often alternate, sometimes opposite, petiolate or sessile, simple, entire, sometimes reduced to scales, exstipulate. Flowers in various sorts of essentially cymose inflorescences, often with a small dichasium axillary to each bract, small, often greenish, hermaphrodite or unisexual (then plants monoecious or dioecious), regular, monochlamydeous, the lobes distinct (in tropical African genera), forming a valvate 3–5-lobed fleshy cup or tube. Stamens as many as and opposite the lobes, inserted at or below their base; anthers (in tropical African genera) with 2 parallel thecae opening by longitudinal slits. Disk epigynous, intrastaminal, lobed, often lining at least the lower part of the perianth tube. Ovary (in tropical African genera) inferior, unilocular; placenta erect, free-central with 2–3 pendulous ovules; style simple, cylindric or sometimes nearly wanting; stigma terminal, capitate or 2–3(–5)-lobed. Fruit indehiscent, dry or fleshy (nut or drupe). Seed solitary; testa obsolete; cotyledons surrounded by copious fleshy, oily or starchy endosperm
Shrubs or trees, root parasites [stem parasites], evergreen or deciduous, synoecious or dioecious [monoecious, andromonoecious]. Leaves opposite [alternate, whorled], simple; stipules absent; petiole present; blade margins entire; venation pinnate. Inflorescences unisexual or bisexual, axillary or terminal, thyrses or umbels [spikes, racemes, cymes, panicles, fascicles], or flowers solitary. Flowers bisexual or unisexual; perianth and androecium perigynous or epigynous; hypanthium adnate to ovary proximally, free distally [completely adnate to ovary]; sepals 0; petals 3–4(–5), distinct, valvate, post-staminal hairs present or absent; nectary present [absent]; stamens 3–4(–5), opposite petals, distinct, free; anthers dehiscing by longitudinal slits; staminodes present in pistillate flowers; pistil 1, 1–3-carpellate, ovary 1/2-inferior or inferior, 1–3-locular proximally, 1-locular distally, placentation free central, pendulous; ovules 2–4 per ovary, anatropous; style 1; stigma 1. Fruits pseudodrupes (mesocarp hard). Seeds 1 per fruit.
Flowers in various sorts of essentially cymose inflorescences, often with a small dichasium axillary to each bract, small, often greenish, hermaphrodite or unisexual (then plants monoecious or dioecious), regular, monochlamydeous, the lobes distinct (in genera of the Flora Zambesiaca area), forming a valvate 3–5-lobed fleshy cup or tube
Ovary (in genera of the Flora Zambesiaca area) inferior, unilocular, placenta erect, free-central with 2–3 pendulous ovules; style simple, cylindric or sometimes nearly wanting; stigma terminal, capitate or 2–3(5)-lobed
Stamens as many as and opposite the lobes, inserted at or below their base; anthers (in genera of the Flora Zambesiaca area) with 2 parallel thecae opening by longitudinal slits
Leaves often alternate, sometimes opposite, petiolate or sessile, simple, entire, sometimes reduced to scales, exstipulate
Stamens the same number as the calyx-lobes and opposite to them; anthers 2-celled, opening lengthwise
Seed solitary; testa obsolete; cotyledons surrounded by copious fleshy, oily or starchy endosperm
Disk epigynous, intrastaminal, lobed, often lining at least the lower part of the perianth tube
Seed without a testa; endosperm copious; embryo straight; cotyledons mostly terete
Calyx often fleshy, adnate to the ovary, lobes 3–6, valvate or slightly imbricate
Leaves alternate or opposite, entire, sometimes reduced to scales, exstipulate
Flowers often greenish, hermaphrodite or unisexual, actinomorphic
Trees, shrubs or herbs, sometimes parasitic on trees or roots
Ovary inferior or half inferior, 1-celled; style simple
Ovules 1–3, pendulous from the top of a basal placenta
Fruit indehiscent, dry or fleshy (nut or drupe)
Herbs, shrubs or trees, hemiparasitic
Fruit nut-like or drupaceous
Disk epigynous
Petals absent
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Foliage retention evergreen
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Hardiness (USDA) 8-11

Usage

Economically, the most important are the fragrant woods of Santalum and Osyris species, particularly Santalum album L. (Indian Sandalwood), S. spicatum (R.Br.) A.DC. (Australian Sandalwood) and Osyris lanceolata Hochst. & Steud. (African Sandalwood) (Christenhusz et al. 2017). Santalum album (Indian Sandalwood) yields an aromatic white timber, and the yellow heartwood and roots are the source of sandal oil used as a body oil, for perfumery and cosmetics as well as a form of incense. Tanning compounds are extracted from Jodina (not in Australia). The fresh leaves of Osyris compressa (P.J.Bergius) A.DC. (syn Colpoon compressum P.J.Bergius) (Cape Sumach, Coast Tannin Bush) in South Africa were used to tan leather a light brown colour, while the bark was used to tan leather dark brown (PlantZAfrica, accessed June 2019). The fruits of a number of species are edible. The fruit of Santalum acuminatum (Sweet Quandong) is used in Australia for jams, jellies and chutneys. The succulent fruiting receptacle of several species of Exocarpos (Ballart), notably E. cupressiformis, is edible, and was used by Australian indigenous peoples and early European settlers. The wood of E. cupressiformis was also used to make spear-throwers as well as bullroarers (Australian Aboriginal Plant Use and Technology pdf). The fruit of Viscum album L. (European Mistletoe, Common Mistletoe), native to Europe, western and southern Asia, is poisonous, and is potentially fatal in a concentrated form. There is extensive mythology and folk medicine based on mistletoes generally and on Viscum album in particular. It is important in northern European culture, folklore and mythology. Species of Viscum and Phoradendron are used as Christmas symbols in Europe and North America respectively (Barlow 1984: 131). Viscum articulatum Burm.f. (Square-stemmed Mistletoe), native to Australia, Papua New Guinea, Malesia to India, has been traditionally used in different parts of the world for treatment of various ailments. It is used in Ayurvedic and Chinese medicine for the management of various diseases (Patel & Singh 2018). Patel and Singh (2018) claim that "modern scientific studies demonstrate efficacy of this plant against hypertension, ulcer, epilepsy, inflammation, wound, nephrotoxicity, HIV, cancer, etc."
Uses medicinal oil poison timber wood
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Cultivation

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