The families of flowering plants.                                                                                                                                                                

Santalaceae R. Br


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IncludingAnthoboleae (Anthobolaceae) Dum., Arionaceae Van Tiegh., Arjonaceae Van Tiegh. corr. Bullock, Canoppiaceae (Canopodaceae) C. Presl, Exocarpaceae Gagnep., Osyrinae (Osyridaceae) Link, Podospermaceae Dulac, Sarcopodaceae Gagnep.Excluding Viscaceae

Habit and leaf form. Trees, shrubs, and herbs. More or less ‘normal’ plants, or switch-plants; sometimes with the principal photosynthesizing function transferred to stems. Leaves well developed, or much reduced. Plants with roots, or rootless (?); partially parasitic. Parasitic on roots of the host, or aerial parts of the host. Stem growth not conspicuously sympodial. Mesophytic, or xerophytic. Leaves opposite (usually), or alternate; ‘herbaceous’, or leathery, or fleshy, or membranous, or modified into spines; petiolate to sessile; gland-dotted, or not gland-dotted; aromatic, or without marked odour; simple; pulvinate. Lamina entire; one-veined, or pinnately veined. Leaves exstipulate.

Leaf anatomy. Stomata present; usually paracytic.

Adaxial hypodermis present, or absent. Lamina dorsiventral, or isobilateral, or centric. Minor leaf veins without phloem transfer cells (Colpoön).

Stem anatomy. Cork cambium present; initially superficial. Nodes unilacunar. Secondary thickening developing from a conventional cambial ring. Xylem with fibre tracheids; with vessels. Vessel end-walls simple. Vessels without vestured pits. Wood parenchyma apotracheal, or paratracheal.

Reproductive type, pollination. Plants hermaphrodite, or monoecious, or dioecious.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes, or in racemes, or in spikes, or in heads, or in fascicles. The ultimate inflorescence unit cymose, or racemose. Inflorescences terminal, or axillary; variously spikes, racemes or heads, the flowers solitary in the bract axils or forming cymes of three. Flowers bracteate; small; regular; (3–)4–5(–8) merous; cyclic; tricyclic. Free hypanthium present, or absent.

Perianth sepaline, or petaline (dubiously, then perhaps sepaline-petaloid); (3–)4–5(–8); free, or joined (the lobes valvate); 1 whorled; sepaloid, or petaloid; green, or white, or cream, or yellow, or red; fleshy, or non-fleshy; persistent, or deciduous. Calyx (‘calycode’) (3–)4–5(–8); 1 whorled; gamosepalous; regular; fleshy, or non-fleshy; persistent, or not persistent; valvate.

Androecium (3–)4–5(–8). Androecial members free of the perianth (attached at its base), or adnate (to the middle of the lobes); all equal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens (3–)4–5(–8); isomerous with the perianth; oppositisepalous. Anthers dorsifixed, or basifixed; dehiscing via pores, or dehiscing via short slits to dehiscing via longitudinal slits; introrse; bilocular, or four locular (Choretrum); tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer; of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate; 3 aperturate; colpate, or porate, or colporate (or colporoidate); 2-celled (in 4 genera).

Gynoecium (2–)3(–5) carpelled. Carpels reduced in number relative to the perianth to isomerous with the perianth. The pistil 1 celled, or (2–)3(–5) celled. Gynoecium syncarpous; synstylovarious, or eu-syncarpous; partly inferior to inferior. Ovary 1 locular (at least above). Epigynous disk commonly present. Gynoecium non-stylate, or non-stylate to stylate, or stylate. Styles 1; apical. Stigmas 1; capitate (or lobed). Placentation basal, or free central. Ovules differentiated to not differentiated; in the single cavity 1–3(–4); pendulous; hemianatropous to anatropous; unitegmic, or without integuments; tenuinucellate. Embryo-sac development Polygonum-type, or Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 1 (trinucleate, Quinchamalium), or 3 (usually); not proliferating; ephemeral. Synergids usually hooked. Endosperm formation cellular, or helobial. Endosperm haustoria present; chalazal, or lateral. Embryogeny variable and often irregular.

Fruit fleshy, or non-fleshy; indehiscent; a drupe, or a nut. The drupes with one stone. Fruit 1 seeded. Seeds endospermic. Endosperm oily, or not oily. Seedswithout a testa. Cotyledons 2. Embryo achlorophyllous (Thesium linifolium).

Seedling.Germination phanerocotylar, or cryptocotylar.

Physiology, biochemistry. Not cyanogenic. Polyacetylenes recorded. Alkaloids present, or absent. Iridoids not detected. Proanthocyanidins present (Gibbs 1974), or absent. Ellagic acid absent (Thesium). Saponins/sapogenins absent. Sugars transported as sucrose (in Santalum). C3. C3 physiology recorded directly in Comandra. Anatomy non-C4 type (Comandra).

Geography, cytology. Temperate to tropical. Cosmopolitan, except in cold regions. X = 5–7, 12, 13 (or more).

Taxonomy.Subclass Dicotyledonae; dubiously Crassinucelli. Dahlgren’s Superorder Santaliflorae; Santalales. Cronquist’s Subclass Rosidae; Santalales. APG 3 core angiosperms; core eudicot; Superorder Santalanae; Order Santalales.

Species 400. Genera 38; Acanthosyris, Amphorogyne, Antholobus, Arjona,Austroamericium, Buckleya, Cervantesia, Choretrum, Cladomyza,Colpoon, Comandra, Daenikera, Dendromyza, Dendrotrophe,Dufrenoya, Elaphanthera, Exocarpos, Geocaulon, Jodina,Kunkeliella, Leptomeria, Mida, Myoschilos, Nanodea,Nestronia, Okoubaka, Omphacomeria, Osyridocarpos, Osyris,Phacellaria, Pyrularia, Quinchamalium, Rhoiacarpos,Santalum, Scleropyrum, Spirogardnera, Thesidium, Thesium.

Economic uses, etc. Some produce edible fruit, and Santalum album is the source of timber and perfume (sandalwood/sandalwood oil).


  • Technical details: Osyris, Santalum. 
  • Technical details: Osyris (Thonner).
  • Technical details: Thesium, Myoschylos. 
  • Thesium humifusum (B. Ent.).
  • Thesium humifusum: Eng. Bot. 1248, 1868.
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