| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Bruniaceae DC.

Including Berzeliaceae Nak.

Habit and leaf form. Ericoid shrubs (usually, or undershrubs), or trees (rarely). Leaves persistent; small; alternate; spiral (usually in five rows); mostly subtrigonous and acerose; imbricate (usually), or not imbricate; non-sheathing; simple. Lamina entire; parallel-veined (usually three veined, occasionally with 5 or up to twenty veins). Leaves exstipulate (or stipules vestigial).

Leaf anatomy. The leaf lamina bifacial to centric (commonly), or dorsiventral (partly or wholly, in a few species). Stomata present; anomocytic. Hairs present; glandular; mostly long, slender and unicellular. Unicellular hairs simple. Complex hairs absent. The mesophyll containing crystals, or without crystals. The crystals when present, druses, or solitary-prismatic.

Axial (stem, wood) anatomy. Cork cambium present; initially superficial. Primary vascular tissues in a cylinder, without separate bundles; collateral. Secondary thickening developing from a conventional cambial ring.

The vessel end-walls oblique; scalariform. The axial xylem with fibre tracheids. The parenchyma scanty paratracheal.

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

Inflorescence, floral, fruit and seed morphology. Flowers solitary (occasionally), or aggregated in ‘inflorescences’ (usually); when solitary, terminal, or axillary; when aggregated, in spikes, or in heads. Inflorescences of sessile flowers, in spikes or more often in heads, the latter sometimes in panicles or racemes, occasionally the flowers solitary and terminal or axillary; with involucral bracts, or without involucral bracts; pseudanthial, or not pseudanthial. Flowers usually 5 bracteate; small (mostly), or medium-sized; regular; (4–)5 merous; cyclic; tetracyclic. Free hypanthium present, or absent. Hypogynous disk present, or absent; when present, intrastaminal.

Perianth with distinct calyx and corolla; (8–)10; 2 whorled; isomerous. Calyx (4–)5; 1 whorled; polysepalous, or gamosepalous; when gamosepalous, (4-)5 blunt-lobed; regular; persistent; imbricate. Corolla (4–)5; 1 whorled; polypetalous, or gamopetalous (in Lonchostoma); imbricate; regular; persistent (often), or deciduous. Petals clawed (often), or sessile.

Androecium (4–)5 (often persistent). Androecial members free of the perianth (usually), or adnate (the filaments sometimes adnate to the claws of the petals to form a tube, or in Lonchostoma the anthers subsessile and borne on the corolla tube); all equal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens (4–)5; isomerous with the perianth; oppositisepalous; alternating with the corolla members; inflexed in bud; almost with sessile anthers (Lonchostoma), or filantherous. Anthers dorsifixed; versatile (often), or non-versatile; dehiscing via longitudinal slits; introrse; appendaged, or unappendaged. The anther appendages when present, apical (by prolongation of the connective). Pollen shed as single grains. Pollen grains aperturate; 3 aperturate, or 6–11 aperturate; colporate (or colporoidate).

Gynoecium 2 carpelled (usually), or 3 carpelled (Audouinia), or 1 carpelled (seemingly, in Mniothamnea and Berzelia). Carpels reduced in number relative to the perianth. The pistil 1–3 celled. Gynoecium monomerous (or pseudomonomerous?), or syncarpous; of one carpel (Mniothamnea, Berzelia), or synovarious to synstylovarious; partly inferior to inferior (usually), or superior (Raspailia). Carpel when seeming monomerous, stylate; apically stigmatic; 1 ovuled. Placentation apical. Ovary when syncarpous, 2 locular (usually), or 3 locular (Audouinia). Gynoecium stylate. Styles when syncarpous, 2(–3); usually partially joined (at least basally); apical. Stigmas 2(–3). Placentation when syncarpous, apical (from near the top of the septum). Ovules 2–4(–12) per locule; funicled; pendulous; epitropous; with ventral raphe; arillate, or non-arillate; anatropous; unitegmic; crassinucellate. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped.

The fruiting carpel when monomerous, indehiscent; an achene, or nucular. Fruit when syncarpous, dehiscent, or indehiscent, or a schizocarp. Mericarps when schizocarpic, 2, or 3; comprising achenes, or comprising follicles, or comprising nutlets. Fruit when syncarpous/non-schizocarpic, a capsule, or achene-like, or a nut; 1 seeded (when nutlike), or 2 seeded (when capsular). Seeds copiously endospermic; very small. Embryo well differentiated. Cotyledons 2. Embryo straight.

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. Iridoids not detected. Proanthocyanidins present. Ellagic acid absent. Aluminium accumulation not found.

Geography, cytology. Cape. Temperate to sub-tropical. South Africa. X = 16 (Staavia).

Taxonomy. Subclass Dicotyledonae; Crassinucelli, or Tenuinucelli (? - embryological and other evidence being conflicting). Dahlgren’s Superorder Rosiflorae; Cunoniales. Cronquist’s Subclass Rosidae; Rosales. APG III core angiosperms; core eudicot; Superorder Asteranae; campanulid. APG IV Order Bruniales.

Species 75. Genera 12; Audouinia, Berzelia, Brunia, Linconia, Lonchostoma, Mniothamnea, Nebelia, Pseudobaeckea, Raspalia, Staavia, Thamnea, Tittmannia.

Illustrations. • Le Maout and Decaisne: Brunia. • Audouinia capitata, Berzelia lanuginosa, Brunia nodiflora (cf. Widdringtonia), Nebelia laevis (as Diberara), Lonchostoma monogynum (as L. monostylis): Engler & Drude, Die Vegetation der Erde 9 (1910). • Brunia microphylla, as Raspalia: Thonner. • Brunia callunoides, as Berzelia: Hook. Ic. Pl. 11 (1867–71). • Brunia phylicoides, as Raspalia passerinoides: Hook. Ic. Pl. 16 (1886). • Staavia dodii: Hook. Ic. Pl. 26 (1898). • Thamnea depressa: Hook. Ic. Pl. 11 (1867–71).

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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