| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Ruscaceae Spreng.

~ Former Liliaceae, Asparagaceae-Nolinoideae of APG III.

Including Phlebaceae Dulac (p.p.).

Habit and leaf form. Shrubs (or subshrubs, often thicket forming), or lianas. Switch-plants; with the principal photosynthesizing function transferred to stems (in the form of glabrous, ovate to lanceolate, alternate, opposite or whorled, pointed phylloclades). Leaves much reduced (to scales). Self supporting, or climbing. Leaves minute; alternate, or opposite, or whorled; membranous.

Leaf anatomy. Stomata anomocytic. The mesophyll containing mucilage cells, or not containing mucilage cells (? — the leaves being reduced, but mucilage cells with raphides common throughout the plants); containing crystals (?), or without crystals. The crystals if present in the reduced leaves, raphides (there being raphide cells throughout the plants). Minor leaf veins without phloem transfer cells (Ruscus).

Axial (stem, wood) anatomy. Secondary thickening absent. The axial xylem with vessels, or without vessels.

The vessel end-walls scalariform.

Root anatomy. Roots with velamen (known in Semele), or without velamen. Root xylem with vessels; vessel end-walls scalariform, or simple.

Reproductive type, pollination. Unisexual flowers present, or absent. Plants hermaphrodite, or monoecious, or andromonoecious (rarely), or dioecious, or polygamomonoecious. Gynoecium of male flowers pistillodial, or vestigial, or absent.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’. Inflorescences in racemelike clusters on the surfaces of the phylloclades (Ruscus), umbel-like clusters on their edges (Semele), or in terminal racemes separate from them (Danaë); racemes, racemelike or umbel-like. Flowers small (inconspicuous, articulated on their pedicels); regular; 3 merous; cyclic. Perigone tube present (urceolate, in Danaë), or absent. Hypogynous disk absent.

Perianth of ‘tepals’; 6; free, or joined; 2 whorled; isomerous; sepaloid, or petaloid; more or less similar in the two whorls, or different in the two whorls (the inner members sometimes narrower); green, or white, or cream; fleshy (sometimes with a fleshy corona), or non-fleshy.

Androecium 3, or 6. Androecial members free of the perianth; coherent (the filaments joined to form a short tube or column); 1 whorled, or 2 whorled. Androecium exclusively of fertile stamens. Stamens 3, or 6; isomerous with the perianth; oppositiperianthial (when 3), or alterniperianthial. Anthers sessile; dehiscing via longitudinal slits; extrorse; tetrasporangiate. Endothecium developing fibrous thickenings. The endothecial thickenings spiral. Microsporogenesis successive. The initial microspore tetrads isobilateral. Tapetum glandular. Pollen shed as single grains. Pollen grains nonaperturate (at least in some species); 2-celled (in Danaë and Ruscus).

Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil 1 celled, or 3 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1 locular, or 3 locular; sessile. Gynoecium shortly stylate. Styles 1; apical. Stylar canal present. Stigmas 1; wet type; papillate; Group III type. Placentation basal; axile. Ovules in the single cavity 2; 2 per locule; collateral; orthotropous, or hemianatropous; bitegmic; tenuinucellate. Embryo-sac development Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (to form 5 cells), or not proliferating; ephemeral. Synergids hooked (with filiform apparatus).

Fruit fleshy; indehiscent; red, a berry; 1–4 seeded. Seeds endospermic. Embryo well differentiated. Cotyledons 1. Embryo achlorophyllous (2/3); cylindrical, short, straight. Testa without phytomelan; pale.

Seedling. Hypocotyl internode present (short). Seedling collar not conspicuous. Coleoptile absent. Seedling cataphylls present. Primary root persistent.

Physiology, phytochemistry. Not cyanogenic. Alkaloids absent (?). Anthraquinones detected (Ruscus); polyacetate derived. Saponins/sapogenins present (steroidal). Proanthocyanidins absent. Flavonols present (Danaë), or absent (Ruscus, Semele); in Danaë, kaempferol and quercetin. Ellagic acid absent.

Geography, cytology. Holarctic, Paleotropical, and Cape. Temperate. Western and Central Europe, Mediterranean.

Taxonomy. Subclass Monocotyledonae. Dahlgren et al. Superorder Liliiflorae; Asparagales. APG III core angiosperms; Superorder Lilianae; non-commelinid Monocot. APG IV Order Asparagales (as a synonym of Asparagaceae).

Species 9. Genera 3; Ruscus, Semele, Danaë.

General remarks. Differing from Asparagaceae sensu stricto (q.v.) in conspicuous features of the androecium, stamen morphology and pollen form, as well as embryology.

Illustrations. • Danaë racemosus (as Ruscus): Baillon, Histoire des Plantes 12 (1894). • Le Maout and Decaisne: Ruscus aculeatus. • Ruscus aculeatus (B. Ent.). • Semele androgyna (as Ruscus): Bot. Mag. 57 (1830).

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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