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Santalaceae R. Brown

檀香科 tan xiang ke

Authors: Nianhe Xia & Michael G. Gilbert

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+.

About 36 genera and 500 species: widely distributed in tropical and temperate regions; seven genera and 33 species (13 endemic) in China.

Three species of sandalwood, Santalum album Linnaeus, S. myrtifolium Linnaeus, and S. papuanum Summerhayes, have been recorded as cultivated in China. Wu Zhengyi (editor’s note) adds that he collected a fruiting specimen of a Santalum in the coastal forests of Taiwan (at KUN).

Tam Pui-cheung. 1988. Santalaceae. In: Kiu Hua-shing & Ling Yeou-ruenn, eds., Fl. Reipubl. Popularis Sin. 24: 52–86.

1 Leaves and vegetative stems almost absent; aerial epiparasite growing on Loranthaceae or Dendrotrophe; flowers minute, sessile, often very densely clustered.   7 Phacellaria
+ Leaves and vegetative stems usually well developed; trees, shrubs, or vines, or terrestrial herbs with scale-like leaves; flowers often large, pedicellate, solitary, paired, or clustered.   (2)
2 (1) Terrestrial herbs; leaves linear or scale-like; each flower subtended by 1, 2, or rarely 4, prominent, persistent bracteoles; fruit a nutlet, exocarp dry [sometimes fleshy].   5 Thesium
+ Trees, shrubs, or vines; leaves well developed, lanceolate to ovate; flowers usually without conspicuous, persistent bracteoles; fruit a drupe, exocarp fleshy.   (3)
3 (2) Leaves opposite, first-formed leaves on shoots with prominent scarious, scale-like tips; fruit with 4(or 5) conspicuous, persistent, accrescent bracts near apex.   1 Buckleya
+ Leaves alternate, tips never scarious; fruit without enlarged persistent bracts.   (4)
4 (3) Leaf blade palmately 3-9(-11)-veined; plants woody vines, often an aerial hemiparasite.   6 Dendrotrophe
+ Leaf blade pinnately veined; plants freestanding trees or shrubs.   (5)
5 (4) Inflorescences cymose or 1-flowered; flowers 3(or 4)-merous; stems usually 3-angled; fruit to 1 cm.   4 Osyris
+ Inflorescences racemelike, thyrsoid, catkinlike, or 1-flowered; flowers 5(or 6)-merous; stems terete; fruit 3-5 cm.   (6)
6 (5) Inflorescences terminal, on leafy axillary shoots, female inflorescences with flowers subtended by well-developed leaves and effectively solitary and axillary; anther cells longitudinally dehiscent; fruit with apical disk more than 5 mm wide.   2 Pyrularia
+ Inflorescences axillary, lacking normal leaves, male inflorescences sometimes in cluster; anther cells transversely dehiscent; fruit with apical disk to 2.5 mm wide.   3 Scleropyrum

Lower Taxa


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