| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Sambucaceae Link.

~ Caprifoliaceae, Adoxaceae sensu lato.

Excluding Viburnaceae Dum.

Habit and leaf form. Small trees, or shrubs, or herbs (few). Plants green and photosynthesizing. Leptocaul. Mesophytic. Leaves deciduous; opposite; petiolate; foetid; compound; pinnate, or pinnate and bipinnate (in that in some species, some leaflets may be paired); imparipinnate. Lamina pinnately veined; cross-venulate. Leaves stipulate (sometimes with four or more stipules side by side at a node), or exstipulate. Stipules when present, often scaly. Lamina margins serrate. Vegetative buds scaly. Leaf development not ‘graminaceous’.

General anatomy. Plants with ‘crystal sand’ (this abundant).

Leaf anatomy. The leaf lamina dorsiventral. Stomata present; mainly confined to one surface (abaxial); mostly anomocytic. Hairs present; eglandular and glandular; unicellular and multicellular (the eglandular ones simple and unicellular, the glandular ones with uniseriate stalk and multicellular, ellipsoidal head). Unicellular hairs simple. Complex hairs absent.

Axial (stem, wood) anatomy. Pith homogeneous (wide, spongy, consisting of dead cells in mature shoots). Cork cambium present; initially superficial. Nodes tri-lacunar, or penta-lacunar, or multilacunar; exhibiting on either side a trace which divides, contributing the outermost lateral traces to each of the opposite leaves (at least in S. nigra and S. ebulus: see illustration), or without split-lateral traces (?). Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring.

The wood semi-ring porous, or diffuse porous. The vessels small; radially paired, in radial multiples, and in tangential arcs. The vessel end-walls simple, or scalariform and simple. The vessels without vestured pits; with spiral thickening (slight), or without spiral thickening. The axial xylem without tracheids; with libriform fibres. The fibres with spiral thickening. The parenchyma scanty paratracheal; wood partially storied (VP). Tyloses present (often abundant), or absent.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in cymes, in corymbs, and in panicles. The ultimate inflorescence units cymose. Inflorescences terminal; large, repeatedly branched, compound, flat topped, umbelliform corymbs and thyrses. Flowers bracteolate (with articulate pedicels); small, or medium-sized; fragrant, or malodorous (a matter of opinion); regular; 3–5 merous; cyclic; tetracyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; 6–10; 2 whorled; isomerous. Calyx 3–5; 1 whorled; gamosepalous; regular; persistent; open in bud; with the median member posterior. Corolla 3–5; 1 whorled; gamopetalous; imbricate, or valvate; rotate (i.e., the tube very short); regular; white.

Androecium 3–5. Androecial members adnate (to the corolla); free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens 3–5; isomerous with the perianth; oppositisepalous; alternating with the corolla members. Anthers dehiscing via longitudinal slits; extrorse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate; 3 aperturate; colporate; 3-celled.

Gynoecium 3–5 carpelled. Carpels reduced in number relative to the perianth to isomerous with the perianth. The pistil 3–5 celled. Gynoecium syncarpous; synstylovarious; inferior. Ovary 3–5 locular. Epigynous disk present, or absent. Gynoecium more or less non-stylate. Stigmas 3–5; dry type; papillate; Group II type. Placentation axile, or apical. Ovules 1 per locule; pendulous; epitropous; with ventral raphe; anatropous; unitegmic; tenuinucellate. Embryo-sac development Adoxa-type (Adoxa type). Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped. Endosperm formation cellular.

Fruit fleshy; indehiscent; a drupe (berrylike). The drupes with separable pyrenes (with 3–5 one-seeded pyrenes). Fruit 3–5 seeded. Seeds endospermic. Endosperm oily. Embryo chlorophyllous (1/1); straight.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Sambucus. Cyanogenic, or not cyanogenic (in different species). Arbutin absent. Iridoids detected; ‘Route I’ type (normal and seco). Proanthocyanidins absent. Flavonols present; kaempferol and quercetin. Ellagic acid absent. Ursolic acid present.

Geography, cytology. Temperate to tropical. Widespread, but absent from Amazonia, Arabia, India, Western Australia and Pacific, and represented in Africa by only one species in the Eastern mountains.

Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Corniflorae; Cornales. Cronquist’s Subclass Asteridae; Dipsacales. APG III core angiosperms; core eudicot; Superorder Asteranae; campanulid. APG IV Order Dipsacales (as a synonym of Adoxaceae).

Species up to 40 (with numerous segregates). Genera 1; only genus, Sambucus.

General remarks. RbcL sequence analyses by Backlund and Bremer (1997) imply close relationship between Adoxa, Sambucus and Viburnum; strongly supporting the taxonomic integrity of Bentham and Hooker’s tribe Caprifoliaceae-Sambuceae (= Adoxaceae sensu lato, e.g. Judd et al. 1984) but suggesting that these genera are relatively distant from Caprifoliaceae. Perhaps their true affinities (a bone of contention since the nineteenth century) have now been resolved. In any case, comparison of this description with that of Caprifoliaceae sensu stricto (q.v.) reveals differences in 17 characters, representing vegetative morphology and anatomy, inflorescence, flowers (perianth, androecium, gynoecium, ovules), embryo .....

Economic uses, etc. Mature berries of some (notably, S. nigra) are used to make wine, and flowers provide flavouring and are used in herbal medicine (e.g., as diuretic). Most have poisonous properties, especially in young shoots, and S. racemosa is very poisonous. 'White elder pith' is much used by plant anatomists to support material for sectioning.

Quotations.

What says my Aesculapius, my Galen, my heart of elder?
(i.e. by contrast with ‘heart-of-oak’: ‘Merry Wives’, ii., 3)

And let the stinking elder, grief, entwine,
His perishing roots with the increasing vine
(‘Cymbeline’, iv., 2 — the foetid S. ebulus)

HOLOFERNES: Begin, sir, you are my elder.
BIRON: Well followed: Judas was hanged on an elder
(‘Love’s Labour’s Lost’, v., 2)

If the medicinal properties of the leaves, bark, berries, etc., were thoroughly known, I cannot tell what our countryman could ail for which he might fetch a remedy from every hedge, either for sickness or wound.
John Evelyn (1664), ‘Sylva’ (of S. nigra).

Illustrations. • Le Maout and Decaisne: Sambucus. • Sambucus nigra: Eng. Bot. 637 (1865). • Sambucus nigra: Köhler's Medizinal Pflanzen 1 (1887). • Sambucus nigra: Lindley. • Sambucus ebulus: Eng. Bot. 638 (1865). • Sambucus ebulus: peculiar nodal anatomy.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 8th June 2026. delta-intkey.com’.

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