BRUNIALES Dumort

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

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.

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.

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

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 = ?

[Audouinieae+Brunieae]

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).

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

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?

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?

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.

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BRUNIALES Bercht. et J. Presl

BRUNIACEAE R. Br. ex DC.

COLUMELLIACEAE D. Don