BRUNIALES Bercht. et J. Presl

Presl, Přir. Rostlin: 226. Jan-Apr 1820 [‘Bruniaceae’]


de Candolle, Prodr. 2: 43. Nov 1825, nom. cons.

Berzeliaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 241. 20 Jul 1943; Brunianae Doweld, Tent. Syst. Plant. Vasc.: liii. 23 Dec 2001

Genera/species 6/81

Distribution Southern coastal and mountain areas in South Africa (almost restricted to Western and Eastern Cape), one species of Brunia in southern KwaZulu-Natal.

Fossils The fossilized flowers of Actinocalyx from the Santonian to the Campanian of Sweden are similar to those in Bruniaceae.

Habit Bisexual, evergreen ericoid shrubs (rarely trees). Lignotubers present or absent.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with scalariform perforation plates; lateral pits scalariform to opposite, simple or bordered pits. Imperforate tracheary xylem element tracheids (in some species of Berzelia and Staavia with crassulae, thickenings of primary walls between tracheids and vessels or among tracheids) with bordered pits, non-septate? Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse (rarely diffuse-in-aggregates; sometimes paratracheal scanty vasicentric). Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Amorphous dark-staining material present in wood of all species. Rhomboidal calciumoxalate crystals usually present (in wood rays, sometimes in axial parenchyma); druses present.

Trichomes Hairs unicellular or multicellular, simple.

Leaves Alternate (spiral, usually pentastichous), simple, entire, usually ericoid, with ? ptyxis. Stipules minute or absent (in Staavia resembling colleters); leaf sheath absent. Venation parallel, usually with three (rarely five to 20) veins, or leaves one-veined. Stomata usually anomocytic (sometimes cyclocytic, with transverse orientation), sometimes on adaxial side of lamina only. Cuticular wax crystalloids as platelets, scales or rods. Mesophyll cells tanniniferous. Rhomboidal crystals frequent; druses present in mesophyll cells in some species. Leaf margin entire, often inrolled. Leaf apex with localized phellogen (cork cells formed from local phellogen), often black, glanduliferous.

Inflorescence Terminal or axillary, usually headlike (sometimes spike-like), these sometimes in panicles or raceme-like partial inflorescences, sometimes (Staavia) in pseudanthia with involucrum consisting of several bracts (in Brunia subgenus Pseudobaeckia solitary along branches; in Thamnea solitary terminal).

Flowers Usually actinomorphic (sometimes slightly zygomorphic), usually small. Usually epigyny or half epigyny (in Brunia subgenus Raspalia hypogyny). Sepals (four or) five, with imbricate aestivation, persistent, free or connate at base. Petals (four or) five, with imbricate aestivation, often clawed, persistent or caducous, free or more or less connate (initially free), in lower part with non-vascularized adaxial ridges or swellings. Nectaries intrastaminal, gynoecial (on upper part of ovary), often as nectariferous disc, or absent.

Androecium Stamens (four or) five, haplostemonous, antesepalous, alternipetalous, often persistent; abaxial stamens larger than the remainder. Filaments free from each other, usually free from petals (sometimes adnate to petals in lower part [epipetalous]). Anthers dorsifixed, inflexed in bud, usually versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective sometimes prolonged at apex. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpor(oid)ate or 6–10(–11)-colpor(oid)ate, shed as monads, ?-cellular at dispersal. Exine tectate or semitectate, with columellate infratectum, imperforate, perforate or reticulate, foveolate, psilate, granulate, regulate, or verrucate.

Gynoecium Pistil composed of usually two (rarely three to five) connate carpels (in Berzelia and Brunia subgenus Mniothamnea at least seemingly one carpel). Ovary usually inferior or semi-inferior, usually bilocular (rarely monomerous or pseudomonomerous? or trilocular to quinquelocular). Stylodia usually two (rarely three to five), almost free to nearly entirely connate (style rarely single, simple). Stigmas capitate, papillate?, Wet type. Pistillodium absent.

Ovules Placentation apical-axile. Ovules (one or) two to four (to twelve) per carpel (when a single carpel, then one ovule), usually anatropous (sometimes pleurotropous), pendulous, epitropous, unitegmic, tenuinucellar to weakly crassinucellar. Micropyle long. Integument massive, ? cell layers thick. Endothelium present. Hypostase present. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A schizocarp usually with two one-seeded nutlike or follicle-like mericarps, a nutlet or a two-seeded loculicidal capsule (sometimes also adaxially dehiscent) with persistent and sometimes accrescent calyx.

Seed: Seed sometimes with aril. Exotesta thin. Endotesta? Perisperm not developed. Endosperm copious, fleshy. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology n = 10, 11, 20–23 – Polyploidy occurring in Berzelia.


Phytochemistry Insufficiently known. Flavonols (quercetin, myricetin, isorhamnetin) and proanthocyanidins (prodelphinidins) present. Ellagic acid not found. Iridoids?

Use Ornamental plants.

Systematics The sister-group relationships of Bruniaceae are not entirely clarified, yet they may be sister to Columelliaceae. Linconia is sister to the remaining Bruniaceae, and the [Audouinia+Thamnea] clade is sister to the clade [Staavia+[Berzelia+Brunia]] (Quint & Claßen-Bockhoff 2006a; Claßen-Bockhoff & al. 2011).

Linconieae Quint et Class.-Bockh. in Taxon 60: 1144. 4 Aug 2011

1/3. Linconia (3; L. alopecuroidea, L. cuspidata, L. ericoides; Western Cape, southwestern Eastern Cape). Lignotubers present. Each stoma surrounded by cuticular rim. Petals stiff and inflexible. Anthers sagittate, with thecae apically connate and with apex ending in sterile conspicuous tip. n = ?


Petals flexible. Anthers without sterile apex.

Audouinieae Nied. in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. III, 2a: 133. 9 Mar 1891

2/14. Audouinia (5; A. capitata, A. esterhuyseniae, A. hispida, A. laevis, A. laxa; Western Cape), Thamnea (9; Western Cape). – Lignotubers present. Stomata often surrounded by cuticular rim. Anthers without sterile apex, with thecae connate along their entire length.Audouinia with thecae entirely adnate to connective and with ovary usually trilocular. n = 11 (Audouinia).

Brunieae Quint et Class.-Bockh. in Taxon 60: 1146. 4 Aug 2011

3/64. Staavia (11; Western Cape), Berzelia (16; Western and Eastern Cape), Brunia (37; Western Cape, southern Eastern Cape, one species in KwaZulu-Natal). – Western and Eastern Cape, KwaZulu-Natal. Lignotubers present or absent. Stomata not surrounded by cuticular rim. Anthers versatile, without sterile apex. Petals and filaments in Brunia subgenus Lonchostoma secondarily postgenitally connate. – Staavia is sister to [Berzelia+Brunia].

Cladogram of Bruniaceae based on DNA sequence data (Quint & Claßen-Bockhoff 2006a).


( Back to Bruniales )

Don in Edinburgh New Philos. J. 6: 46, 49. Oct-Dec 1828 [’Columellieae’], nom. cons.

Columelliales D. Don in C. F. P. von Martius, Consp. Regn. Veg.: 27. Sep-Oct 1835 [’Columellieae’]; Desfontainiaceae (Endl.) L. K. G. Pfeiffer, Nomencl. Bot. 1: 1037. 21 Feb 1873 [’Desfontaineae’], nom. cons. Desfontainiales Takht., Divers. Classif. Fl. Pl.: 377. 24 Apr 1997

Genera/species 2/8

Distribution Southern Mexico to Cape Horn in Chile.

Fossils Unknown.

Habit Bisexual, evergreen shrubs or small trees. Bud scales absent.

Vegetative anatomy Phellogen ab initio pericyclic (in Desfontainia near phloem). Primary medullary rays in Desfontainia very thin, mostly uniseriate. Medulla parenchymatous (Desfontainia). Pericyclic fibres absent. Vessel elements with scalariform perforation plates; lateral pits scalariform (Desfontainia), or alternate or absent (Columellia), simple or bordered pits. Vestured pits absent. Imperforate tracheary xylem elements tracheids or fibre tracheids with simple or bordered pits, non-septate (also vasicentric or vascular tracheids). Wood rays uniseriate, heterocellular. Axial parenchyma apotracheal usually diffuse, or paratracheal vasicentric (Columellia). Intraxylary phloem absent. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace (Columellia). Pericycle with scattered sclereids (Desfontainia). Calciumoxalate as druses.

Trichomes Hairs simple (sometimes absent in Desfontainia); glands and glandular hairs present or absent.

Leaves Opposite, simple, entire, in Columellia strongly asymmetrical, in Desfontainia coriaceous, with conduplicate ptyxis. Stipules small interpetiolar to intrapetiolar (Desfontainia) or absent (Columellia, sometimes in Desfontainia); leaf sheath absent. Colleters present in Desfontainia. Petiole vascular bundle transection arcuate; central vascular bundle largest (Columellia). Abaxial side of lamina in Columellia sometimes glanduliferous. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids in at least Columellia as tubuli (scarcely branched), dominated by β-diketones. Mesophyll in Columellia with calciumoxalate druses. Leaf margin entire to serrate, usually with glandular tip (Columellia) or serrate-dentate to spinose-serrate (Desfontainia).

Inflorescence Terminal? (Columellia) or axillary? (Desfontainia), few-flowered cymes or flowers solitary (in Desfontainia usually solitary).

Flowers Actinomorphic (Desfontainia) or somewhat zygomorphic (Columellia, Desfontainia), medium-sized. Epigyny (Columellia) or hypogyny (Desfontainia). Sepals (four or) five (to eight), with valvate or somewhat imbricate aestivation, persistent, usually more or less connate in lower part; median sepal abaxial. Petals (four or) five (to eight), with imbricate (Columellia) or imbricate to contorted (Desfontainia) aestivation, connate at base (Columellia) or connate (petals possibly initially free), tubular and fleshy (Desfontainia). Nectary absent. Disc absent.

Androecium Stamens in Columellia two (adaxial pair), alternating with adaxial and lateral calyx lobes, alternipetalous; in Desfontainia five, antesepalous, alternipetalous. Filaments short, stout, free, in Columellia inserted at petal base, in Desfontainia adnate to uppermost part of corolla tube. Anthers dorsifixed (Columellia) or basifixed (Desfontainia), non-versatile, tetrasporangiate, latrorse to extrorse (Columellia) or introrse (Desfontainia), longicidal (dehiscing by longitudinal slits); thecae semicircular and attached their entire length to expanded connective (Columellia), or embedded in connective (Desfontainia); connective well developed, in Columellia expanded at apex. Tapetum amoeboid-periplasmodial (Columellia) or secretory (Desfontainia). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, tricellular at dispersal (Columellia). Exine tectate or semitectate, with columellate infratectum, finely reticulate (Columellia) or perforate (Desfontainia), rugulate.

Gynoecium Pistil composed of two median (Columellia) or (three to) five (to seven) alternisepalous, antepetalous (Desfontainia) connate carpels. Ovary inferior (Columellia) or superior (Desfontainia), incompletely bilocular (Columellia), or bilocular to quinquelocular at base and unilocular at apex (Desfontainia). Style single, simple, short and thick (Columellia), or longer and filiform, persistent (Desfontainia). Stigma almost cup-shaped, bilobate or quadrilobate (Columellia), or capitate to somewhat quinquelobate (Desfontainia), type? Pistillodium absent.

Ovules Placentation intrusively parietal to axile; placentae in Columellia forming secondary incomplete septa; placentation in Desfontainia mainly axile yet strongly intrusively parietal in uppermost part. Ovules numerous per carpel, anatropous, ascending (Columellia), unitegmic, tenuinucellar (Columellia) or crassinucellar (Desfontainia). Integument six to nine cell layers thick (Desfontainia). Parietal tissue possibly absent (Columellia). Haustorial suspensor large in Desfontainia. Endothelium poorly developed (Desfontainia). Megagametophyte monosporous, Polygonum type. Endosperm development in Columellia ?, in Desfontainia cellular. Endosperm haustoria in Columellia ?, in Desfontainia chalazal and micropylar. Embryogenesis solanad (Desfontainia).

Fruit A septicidal and partially loculicidal capsule (Columellia) or a berry (Desfontainia) with numerous seeds and persistent calyx.

Seeds Aril? Seed coat exotestal. Exotestal cells elongate (Columellia) or polygonal (Desfontainia); anticlinal exotestal cells strongly thickened, with plasmodesmata; outer walls of exotestal cells pectic and remaining cell walls lignified (Desfontainia). Endotesta? Perisperm not developed. Endosperm copious (in Desfontainia starchy). Embryo small, straight, well differentiated, without chlorophyll. Cotyledons two. Germination?

Cytology n = 7 (Desfontainia)


Phytochemistry Insufficiently known. Route I iridoids (secoiridoids: loganin, loganic acid, etc.) and cornoside present in Desfontainia. Proanthocyanidins, saponins and cyanogenic compounds not found. Tannins not found in Desfontainia. Myricetin?

Use Medicinal plants, carpentry, dyeing substances.

Systematics Columellia (5; C. lucida, C. oblonga, C. obovata, C. subsessilis, C. weberbaueri; the Andes from southern Colombia to Bolivia), Desfontainia (3; D. fulgens: Chile, western Argentina; D. spinosa: the Andes from Costa Rica to Cape Horn in Chile; D. splendens: mountains from southern Mexico to Bolivia).

The sister-group relationship of Columelliaceae is not entirely clarified, although they seem to be sister to Bruniaceae.


Bremer B, Olmstead RG, Struwe L, Sweere JA. 1994. rbcL sequences support exclusion of Retzia, Desfontainia, and Nicodemia from the Gentianales. – Plant Syst. Evol. 190: 213-230.

Brizicky GK. 1961. A synopsis of the genus Columellia (Columelliaceae). – J. Arnold Arbor. 42: 363-372.

Carlquist SJ. 1978. Wood anatomy of Bruniaceae: correlations with ecology, phylogeny, and organography. – Aliso 9: 323-364.

Carlquist SJ. 1990. Leaf anatomy of Geissolomataceae and Myrothamnaceae as a possible indicator of relationship to Bruniaceae. – Bull. Torrey Bot. Club 117: 420-428.

Carlquist SJ. 1991. Leaf anatomy of Bruniaceae: ecological, systematic, and phylogenetic aspects. – Bot. J. Linn. Soc. 107: 1-34.

Claßen-Bockhoff R. 2000. Inflorescences in Bruniaceae, with general comments on inflorescences in woody plants. – Opera Bot. Belg. 12: 5-310.

Claßen-Bockhoff R, Oliver EGH, Hall AV, Quint M. 2011. A new classification of the South African endemic family Bruniaceae based on molecular and morphological data. – Taxon 60: 1138-1155.

Dahlgren RMT, Wyk AE van. 1988. Structures and relationships of families endemic to or centered in southern Africa. – Monogr. Syst. Bot. Missouri Bot. Gard. 25: 1-94.

Daniel M, Sabnis SD. 1979. Chemotaxonomy of Loganiaceae. – Curr. Sci. 48: 383-385.

Dümmer RA. 1912. An enumeration of the Bruniaceae. – J. Bot. 50 [Suppl. 2]: 1-37.

Fagerström K. 1975. 182. Columelliaceae. – In: Harling G, Sparre B (eds), Flora of Ecuador 4, Swedish Natural Science Research Council, Stockholm, pp. 3-5.

Fritsch K. 1895. Columelliaceae. – In: Engler A, Prantl K (eds), Die natürlichen Pflanzenfamilien IV(3b), W. Engelmann, Leipzig, pp. 186-188.

Goldblatt P. 1981. Chromosome cytology of Bruniaceae. – Ann. Missouri Bot. Gard. 68: 546-550.

Gregory M. 1998. Columelliaceae. – In: Cutler DF, Gregory M (eds), Anatomy of the dicotyledons, 2nd ed., IV, Clarendon Press, Oxford, pp. 117-120.

Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related familes (Asterales). – Amer. J. Bot. 82: 250-265.

Hall AV. 1987. Evidence of a Cretaceous alliance for the Bruniaceae. – South Afr. J. Sci. 83: 58-59.

Hall AV. 1988. Systematic palynology of the Bruniaceae. – Bot. J. Linn. Soc. 96: 285-296.

Hall AV, Oliver EGH, Claßen-Bockhoff R. 2010. Bruniaceae: new species of Thamnea and Brunia from western Cape, South Africa. – Bothalia 40: 96-101.

Hasselberg GBE. 1937. Zur Morphologie des vegetativen Sproßes der Loganiaceen. – Symb. Bot. Upsal. 2(3): 1-170.

Hoc P, Bravo L. 1984. Estudio palinológico sobre las especies presentes en Argentina de Spigelia, Strychnos, y Desfontainia (Loganiaceae). – Kurtziana 17: 71-89.

Houghton PJ, Ming L-L. 1985. Iridoids from Desfontainia spinosa. – Phytochemistry 24: 1841-1842.

Houghton PJ, Ming L-L. 1986a. Iridoids, iridoid-triterpenoid congeners and lignans from Desfontainia spinosa. – Phytochemistry 25: 1907-1912.

Houghton PJ, Ming L-L. 1986b. Triterpenoids from Desfontainia spinosa. – Phytochemistry 25: 1939-1944.

Jay MM. 1968. Distribution des flavonoïdes chez les Bruniacées. – Taxon 17: 484-488.

Kadereit JW, Bittrich V (eds). 2016. The families and genera of vascular plants XIV. Flowering plants – eudicots – Aquifoliales, Boraginales, Bruniales, Dipsacales, Escalloniales, Garryales, Paracryphiales, Solanales (except Convolvulaceae), Icacinaceae, Metteniusaceae, Vahliaceae. – Springer, 412 pp.

Kirchner R. 1904. Beiträge zur Kenntniss der Bruniaceen. – Ph.D. diss., Universität Breslau, Poland.

Lange JHD, Boucher C, Walt JJA van der. 1993. Autecological studies on Audouinia capitata (Bruniaceae) 3. Pollination biology. – South Afr. J. Bot. 59: 135-144.

Lange JHD, Walt JJA van der, Boucher C. 1993a. Autecological studies on Audouinia capitata (Bruniaceae) 5. Seed development, abortion and pre-emergent reproductive success. – South Afr. J. Bot. 59: 156-167.

Lange JHD, Walt JJA van der, Boucher C. 1993b. Autecological studies on Audouinia capitata (Bruniaceae) 6. Nutritional aspects of the developing ovule. – South Afr. J. Bot. 59: 168-177.

Leeuwenberg AJM. 1969. Notes on American Loganiaceae IV. Revision of Desfontainia Ruiz et Pav. – Acta Bot. Neerl. 18: 669-679.

Leeuwenberg AJM (ed). 1980. Angiospermae: Ordnung Gentianales Fam. Loganiaceae. – In: Hiepko P, Melchior H (eds), Die natürlichen Pflanzenfamilien, 2. Aufl., Vol. 28bI, Duncker & Humblot, Berlin, pp. 1-255.

Leinfellner W. 1964a. Über die falsche Sympetalie bei Lonchostoma und anderen Gattungen der Bruniaceen. – Österr. Bot. Zeitschr. 111: 345-353.

Leinfellner W. 1964b. Sind die Kronblätter der Bruniaceae peltat gebaut? – Österr. Bot. Zeitschr. 111: 500-526.

Maldonado de Magnano S. 1986. Estudios embriológicos en Desfontainia spinosa (Desfontainiaceae). – Darwiniana 27: 207-224.

Niedenzu F. 1891. Bruniaceae. – In: Engler A, Prantl K (eds), Die natürlichen Pflanzenfamilien III(2a), W. Engelmann, Leipzig, pp. 131-136.

Niedenzu F, Harms H. 1930. Bruniaceae. – In: Engler A, Harms H (eds), Die natürlichen Pflanzenfamilien, 2. Aufl., Bd. 18a, W. Engelmann, Leipzig, pp. 288-303.

Oliver EGH, Oliver IM. 1999. Bruniaceae. A new species of Linconia from Western Cape. – Bothalia 29: 256-258.

Pillans NS. 1947. A revision of Bruniaceae. – J. South Afr. Bot. 13: 121-206.

Powrie E. 1969a. Types of Bruniaceae in the Thunberg Herbarium. – J. South Afr. Bot. 35: 327-339.

Powrie E. 1969b. A new species of Tittmannia (Bruniaceae). – J. South Afr. Bot. 35: 363-366.

Quint M. 2004. Evolution of Bruniaceae: evidence from molecular and morphological studies. – Ph.D. diss., Universität Mainz, Germany.

Quint M, Claßen-Bockhoff R. 2006a. Phylogeny of Bruniaceae based on matK and ITS sequence data. – Intern. J. Plant Sci. 167: 135-146.

Quint M, Claßen-Bockhoff R. 2006b. Floral ontogeny, petal diversity and nectary uniformity in Bruniaceae. – Bot. J. Linn. Soc. 150: 459-477.

Quint M, Claßen-Bockhoff R. 2008. Ancient or recent? Insights into the temporal evolution of the Bruniaceae. – Organisms Divers. Evol. 8: 293-304.

Saxton WT. 1910. The ovule of the Bruniaceae. – Trans. Roy. Soc. South Africa 2: 27-31.

Schlechter R. 1920. Die Columelliaceae. – Notizbl. Bot. Gart. Berlin-Dahlem 7: 352-358.

Schultes RE. 1977. De plantes toxicariis e mundo novo tropicale commentationes: 15. Desfontainia: a new Andean hallucinogen. – Bot. Mus. Leafl. 25: 99-104.

Schultes RE. 1989. De speciebus varietatibusque Desfontainia Columbianae notae. – Rev. Acad. Colomb. Ci. Exact. Fisic. Natur. 17: 313-319.

Scott G. 1999. A chemosystematic and cladistic study of the Southern African endemic family Bruniaceae. – Ph.D. diss., Department of Botany, University of Cape Town, Republic of South Africa.

Solereder H. 1895. Loganiaceae. – In: Engler A, Prantl K (eds), Die natürlichen Pflanzenfamilien IV(2), W. Engelmann, Leipzig, pp. 19-50.

Stern WL, Brizicky GK, Eyde RH. 1969. Comparative anatomy and relationships of Columelliaceae. – J. Arnold Arbor. 50: 36-75.

Strid A. 1968. A new species of Lonchostoma (Bruniaceae). – Bot. Not. 121: 312-316.

Tieghem P van. 1903. Sur les Columelliacées. – Ann. Sci. Nat. 17-18: 155-164.

Villiers SE de, Cadman A. 1997. The palynology of tertiary sediments from a palaeochannel in Namaqualand, South Africa. – Palaeontol. Afr. 34: 69-99.

Weberling F. 1976. Weitere Untersuchungen zur Morphologie des Unterblattes bei den Dikotylen IX. Saxifragaceae s.l., Brunelliaceae, und Bruniaceae. – Beitr. Biol. Pflanzen 52: 163-181.

Weigend M. 2001. Desfontainia Ruiz & Pav. (Desfontainiaceae) revisited – a first step back towards α-diversity. – Bot. Jahrb. Syst. 123: 281-301.