EMBRYOPSIDA Pirani & Prado

Gametophyte dominant, independent, multicellular, thalloid, with single-celled apical meristem, showing gravitropism; rhizoids +, unicellular; acquisition of phenylalanine lysase [PAL], phenylpropanoid metabolism [lignans +, flavonoids + (absorbtion of UV radiation)], xyloglucans +; plant [protoplasm dessication tolerant], ectohydrous [free water outside plant physiologically important]; cuticle +; cell wall also with (1->3),(1->4)-ß-D-MLGs [Mixed-Linkage Glucans]; chloroplasts per cell, lacking pyrenoids; glycolate metabolism in leaf peroxisomes [glyoxysomes]; centrioles in vegetative cells 0, metaphase spindle anastral, predictive preprophase band of microtubules, phragmoplast + [cell wall deposition spreading from around the spindle fibres], plasmodesmata +; antheridia and archegonia jacketed, stalked; spermatogenous cells monoplastidic; blepharoplast, bicentriole pair develops de novo in spermatogenous cell, associated with basal bodies of cilia [= flagellum], multilayered structure [4 layers: L1, L4, tubules; L2, L3, short vertical lamellae] + spline [tubules from L1 encircling spermatid], basal body 200-250 nm long, associated with amorphous electron-dense material, microtubules in basal end lacking symmetry, stellate array of filaments in transition zone extended, axonemal cap 0 [microtubules disorganized at apex of cilium]; male gametes [spermatozoids] with a left-handed coil, cilia 2, lateral; oogamy; sporophyte dependent on gametophyte, embryo initially surrounded by haploid gametophytic tissue, plane of first division horizontal [with respect to long axis of archegonium/embryo sac], suspensor/foot +, cell walls with nacreous thickenings; sporophyte multicellular, with at least transient apical cell [?level], sporangium +, single, dehiscence longitudinal; meiosis sporic, monoplastidic, microtubule organizing centre associated with plastid, cytokinesis simultaneous, preceding nuclear division, sporocytes 4-lobed, with a quadripolar microtubule system; spores in tetrads, sporopollenin in the spore wall, initially laid down in association with several trilamellar layers [white-line centred layers, i.e. walls multilamellate]; nuclear genome size <1.4 pg, LEAFY gene present, ethylene involved in cell elongation; chloroplast genome with close association between trnLUAA and trnFGAA genes.

Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

STOMATOPHYTES

Abscisic acid, ?D-methionine +; sporangium with tapetal layer, columella + [developing from endothecial cells], seta developing from basal meristem [between epibasal and hypobasal cells; stomata +, anomocytic, cell lineage that produces them with symmetric divisions [perigenous]; underlying similarities in the development of conducting tissue and in rhizoids/root hairs; spores trilete; polar transport of auxins and class 1 KNOX genes expressed in the sporangium alone; shoot meristem patterning gene families expressed; MIKC, MI*K*C* and class 1 and 2 KNOX genes, post-transcriptional editing of chloroplast genes; gain of three group II mitochondrial introns.

[Anthocerophyta + Polysporangiophyta]: archegonia embedded/sunken in the gametophyte; sporophyte long-lived, chlorophyllous; sporophyte-gametophyte junction interdigitate, sporophyte cells showing rhizoid-like behaviour.

POLYSPORANGIOPHYTA†

Sporophyte branched, branching apical, dichotomous; sporangia several, each opening independently; spore walls not multilamellate [?here].

EXTANT TRACHEOPHYTA / VASCULAR PLANTS

Photosynthetic red light response; plant homoiohydrous [water content of protoplasm relatively stable]; control of leaf hydration passive; (condensed or nonhydrolyzable tannins/proanthocyanidins +); sporophyte soon independent, dominant, with basipetal polar auxin transport; lignins +; vascular tissue +sieve cells + [nucleus degenerating], tracheids +, in both protoxylem and metaxylem, plant endohydrous [physiologically important free water inside plant]; endodermis +; stem with an apical cell; branching dichotomous; leaves spirally arranged, blades with mean venation density 1.8 mm/mm2 [to 5 mm/mm2]; sporangia adaxial on the sporophyll, derived from periclinal divisions of several epidermal cells, wall multilayered [eusporangium]; columella 0; tapetum glandular; gametophytes exosporic, green, photosynthetic; basal body 350-550 nm long, stellate array in transition region initially joining microtubule triplets; placenta with single layer of transfer cells in both sporophytic and gametophytic generations, root lateral with respect to the longitudinal axis of the embryo [plant homorhizic].

[MONILOPHYTA + LIGNOPHYTA]

Sporophyte branching ± indeterminate; root apex multicellular, root cap +, lateral roots +, endogenous; endomycorrhizal associations + [with Glomeromycota]; G-type tracheids +, with scalariform-bordered pits; leaves with apical/marginal growth, venation development basipetal, growth determinate; sporangia borne in pairs and grouped in terminal trusses, dehiscence longitudinal, a single slit; cells polyplastidic, microtubule organizing centres not associated with plastids, diffuse, perinuclear; blepharoplasts +, paired, with electron-dense material, centrioles on periphery, male gametes multiciliate; chloroplast long single copy ca 30kb inversion [from psbM to ycf2]; LITTLE ZIPPER proteins.

LIGNOPHYTA†

Sporophyte woody; lateral root origin from the pericycle; branching lateral, meristems axillary; cork cambium + [producing cork abaxially], vascular cambium bifacial [producing phloem abaxially and xylem adaxially].

SEED PLANTS†

Plants heterosporous; megasporangium surrounded by cupule [i.e. ovule +, cupule = integument]; pollen lands on ovule; megaspore germination endosporic [female gametophyte initially retained on the plant].

EXTANT SEED PLANTS / SPERMATOPHYTA

Plant evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins particularly with guaiacyl and p-hydroxyphenyl [G + H] units [sinapyl units uncommon, no Maüle reaction]; root stele with xylem and phloem originating on alternate radii, cork cambium deep seated; shoot apical meristem interface specific plasmodesmatal network; stem with vascular cylinder around central pith [eustele], phloem abaxial [ectophloic], endodermis 0, xylem endarch [development centrifugal]; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, sieve tube plastids with starch grains; phloem fibres +; cork cambium superficial; leaves with single trace from vascular sympodium [nodes 1:1]; stomatal pore with active opening in response to leaf hydration, control by abscisic acid, metabolic regulation of water use efficiency, etc.; axillary buds exogenous, (none; not associated with all leaves); prophylls two, lateral; leaves with petiole and lamina, development basipetal, blade simple; plant heterosporous, sporangia borne on sporophylls, sporophylls spiral; microsporophylls aggregated in indeterminate cones/strobili; grains monosulcate, aperture in ana- position [distal], primexine + [involved in exine pattern formation and deposition of sporopollenin from tapetum], exine and intine homogeneous; megasporangium indehiscent; ovules unitegmic, parietal tissue 2+ cells across, megaspore tetrad linear, functional megaspore single, chalazal, sporopollenin 0; gametophytes dependent on sporophyte, apical cell 0, rhizoids 0; male gametophyte development initially endosporic, tube developing from distal end of grain, gametes two, developing after pollination, with cell walls; female gametophyte initially syncytial, walls then surrounding individual nuclei; embryo cellular ab initio, endoscopic, plane of first cleavage of zygote transverse, suspensor +, short-minute, embryonic axis straight [shoot and root at opposite ends; plant allorhizic], cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, whole nuclear genome duplication [zeta duplication], two copies of LEAFY gene, PHY gene duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.

ANGIOSPERMAE / MAGNOLIOPHYTA

Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], lignin also with syringyl units common [G + S lignin, positive Maüle reaction - syringyl:guaiacyl ratio more than 2-2.5:1], hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0, hypodermis suberised and with Casparian strip [= exodermis +]; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, associated gelatinous fibres [g-fibres] with innermost layer of secondary cell wall rich in cellulose and poor in lignin; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma +; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cell and sieve tube from same mother cell; sugar transport in phloem passive; nodes 1:?; stomata brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule, reduction in stomatal conductance to increasing CO2 concentration; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, overall growth ± diffuse, venation hierarchical-reticulate, secondary veins pinnate, veins (1.7-)4.1(-5.7) mm/mm2, endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, ± haplomorphic; protogynous; parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P +, members each with a single trace, outer members not sharply differentiated from the others, not enclosing the floral bud; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], sporangium pairs dehiscing longitudinally by a common slit, ± embedded in the filament, walls with at least outer secondary parietal cells dividing, endothecium +, endothecial cells elongated at right angles to long axis of anther; (tapetum glandular), cells binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellate, endexine lamellate only in the apertural regions, thin, compact; nectary 0; carpels present, superior, free, several, ascidiate, with postgenital occlusion by secretion, stylulus at most short [shorter than ovary], hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, stigma wet, extragynoecial compitum +; ovules few [?1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, functional megaspore, chalazal, lacking cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; ovule not increasing in size between pollination and fertilization; pollen grains land on stigma, bicellular at dispersal, mature male gametophyte tricellular, germinating in less than 3 hours, pollen tube elongated, unbranched, growing between cells, growth rate (20-)80-20,000 µm/hour, apex of pectins, wall with callose, lumen with callose plugs, penetration of ovules via micropyle [porogamous], whole process takes ca 18 hours, distance to first ovule 1.1-2.1 mm; male gametes lacking cell walls, cilia 0, siphonogamy; double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; mature seed much larger than ovule when fertilized, small [], dry [no sarcotesta], exotestal; endosperm diploid, cellular, heteropolar [micropylar and chalazal domains develop differently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], copious, oily and/or proteinaceous; dark reversal Pfr → Pr; Arabidopsis-type telomeres [(TTTAGGG)n]; nuclear genome size <1.4 pg [1 pg = 109 base pairs], whole nuclear genome duplication [epsilon duplication]; protoplasm dessication tolerant [plant poikilohydric]; ndhB gene 21 codons enlarged at the 5' end, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and three copies of the PHY gene, [PHYB [PHYA + PHYC]].

[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: wood fibres +; axial parenchyma diffuse or diffuse-in-aggregates; pollen monosulcate [anasulcate], tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

[AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessel elements with scalariform perforation plates in primary xylem; essential oils in specialized cells [lamina and P ± pellucid-punctate]; tension wood +; tectum reticulate; anther wall with outer secondary parietal cell layer dividing; carpels plicate; nucellar cap + [character lost where in eudicots?]; 12BP [4 amino acids] deletion in P1 gene.

[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]] / MESANGIOSPERMAE: benzylisoquinoline alkaloids +; sesquiterpene synthase subfamily a [TPS-a] [?level], polyacetate derived anthraquinones + [?level]; outer epidermal walls of root elongation zone with cellulose fibrils oriented transverse to root axis; P more or less whorled, 3-merous [possible position]; pollen tube growth intra-gynoecial [extragynoecial compitum 0]; embryo sac bipolar, 8 nucleate, antipodal cells persisting; endosperm triploid.

[MONOCOTS [CERATOPHYLLALES + EUDICOTS]]: (extra-floral nectaries +); (veins in lamina often 7-17 mm/mm2 or more [mean for eudicots 8.0]); (stamens opposite [two whorls of] P); (pollen tube growth fast).

[CERATOPHYLLALES + EUDICOTS]: ethereal oils 0.

EUDICOTS: (Myricetin, delphinidin +), asarone 0 [unknown in some groups, + in some asterids]; root epidermis derived from root cap [?Buxaceae, etc.]; (vessel elements with simple perforation plates in primary xylem); nodes 3:3; stomata anomocytic; flowers (dimerous), cyclic; protandry common; K/outer P members with three traces, ("C" +, with a single trace); A few, (polyandry widespread, initial primordia 5, 10, or ring, ± centrifugal), filaments fairly slender, anthers basifixed; microsporogenesis simultaneous, pollen tricolpate, apertures in pairs at six points of the young tetrad [Fischer's rule], cleavage centripetal, wall with endexine; G with complete postgenital fusion, stylulus/style solid [?here]; seed coat?

[PROTEALES [TROCHODENDRALES [BUXALES + CORE EUDICOTS]]]: (axial/receptacular nectary +).

[TROCHODENDRALES [BUXALES + CORE EUDICOTS]]: benzylisoquinoline alkaloids 0; euAP3 + TM6 genes [duplication of paleoAP3 gene: B class], mitochondrial rps2 gene lost.

[BUXALES + CORE EUDICOTS]: ?

CORE EUDICOTS / GUNNERIDAE: (ellagic and gallic acids +); leaf margins serrate; compitum + [one place]; micropyle?; whole nuclear genome duplication [palaeohexaploidy, gamma triplication], PI-dB motif +, small deletion in the 18S ribosomal DNA common.

[ROSIDS ET AL. + ASTERIDS ET AL.] / PENTAPETALAE: root apical meristem closed; (cyanogenesis also via [iso]leucine, valine and phenylalanine pathways); flowers rather stereotyped: 5-merous, parts whorled; P = calyx + corolla, the calyx enclosing the flower in bud, sepals with three or more traces, petals with a single trace; stamens = 2x K/C, in two whorls, internal/adaxial to the corolla whorl, alternating, (numerous, but then usually fasciculate and/or centrifugal); pollen tricolporate; G [5], G [3] also common, when [G 2], carpels superposed, compitum +, placentation axile, style +, stigma not decurrent; endosperm nuclear; fruit dry, dehiscent, loculicidal [when a capsule]; RNase-based gametophytic incompatibility system present; floral nectaries with CRABSCLAW expression; (monosymmetric flowers with adaxial/dorsal CYC expression).

[SANTALALES [BERBERIDOPSIDALES [CARYOPHYLLALES + ASTERIDS]]] / ASTERIDS ET AL. / SUPERASTERIDS : ?

[BERBERIDOPSIDALES [CARYOPHYLLALES + ASTERIDS]]: ?

[CARYOPHYLLALES + ASTERIDS]: seed exotestal; embryo long.

ASTERIDS / ASTERIDAE / ASTERANAE Takhtajan: nicotinic acid metabolised to its arabinosides; (iridoids +); tension wood decidedly uncommon; C enclosing A and G in bud, (connate [sometimes evident only early in development, petals then appearing to be free]); anthers dorsifixed?; (nectary gynoecial); G [2], style single, long; ovules unitegmic, integument thick, endothelium +, nucellar epidermis does not persist; exotestal [!: even when a single integument] cells lignified, esp. on anticlinal and/or inner periclinal walls; endosperm cellular.

[ERICALES [ASTERID I + ASTERID II]]: (ovules lacking parietal tissue) [tenuinucellate].

[ASTERID I + ASTERID II] / CORE ASTERIDS: ellagic acid 0, non-hydrolysable tannins not common; sugar transport in phloem active; inflorescence basically cymose; A = and opposite sepals or P, (numerous, usu. associated with increased numbers of C or G); style short[?]; duplication of the PI gene.

ASTERID II / CAMPANULIDAE: myricetin 0; vessel elements with scalariform perforation plates; endosperm copious, embryo short/very short.

[ASTERALES [ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]] / APIIDAE: iridoids +; C forming a distinct tube, tube initiation early; A epipetalous; ovary inferior, [2-3], style long[?].

[ESCALLONIALES [BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]]: ?

[BRUNIALES [APIALES [PARACRYPHIALES + DIPSACALES]]]: ?

Age. Magallón and Castillo (2009: but topology) offer estimates of ca 93.75 m.y. for the crown group, while ages in Beaulieu et al. (2013a) are ca 93 m.y. and in Magallón et al. (2015) ca 86.8 m.y. old. Note that many estimates of stem-group ages for Bruniales - implicitly the age of this node - are based on very different topologies that that here, and/or ignore the existence of Columelliaceae as sister to Bruniaceae.

Evolution. Divergence & Distribution. Diversification at this node probably occurred in the southern hemisphere (Beaulieu et al. 2013a). Bruniales are another old, species poor but morphologically quite diverse clade that is placed along the spine of the campanulids.

Phylogeny. For the relationships of Bruniales, see the asterid II clade.

BRUNIALES Dumortier  Main Tree.

Plant woody, evergreen; iridoids?; nodes 1:1; flowers polysymmetric; anthers basifixed; G position?; ovules weakly crassinucellate. - 2 families, 14 genera, 79 species.

Age. Bruniales crown group age is around 110 m.y. (Bremer et al. 2004) or ca 78.3 m.y. (Magallón et al. 2015).

Note: (....) denotes a feature common in the clade, exact status uncertain, [....] includes explanatory material. Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed (see above).

Phylogeny. Within Bruniales, Bruniaceae are sister (1.0 Bayesian p.p.) to Columelliaceae in many analyses in Winkworth et al. (2008a), although not when coding chloroplast sequences were used alone, while Soltis et al. (2011) found strong support for this clade only when mitochondrial genes were removed from the analysis. Both palynology and wood anatomy had suggested a relationship between Columellia and Desfontainia (ex Loganiaceae) (Mennenga, in Leeuwenberg 1980), and this was strongly supported by rbcL and other data (see B. Bremer et al. 1994; Backlund & Bremer 1997; Bell et al. 2001).

Chemistry, Morphology, etc. For more on the inferior/superior ovary distinction in the campanulids, see the Asterales page.

Classification. Allthough Columellia and Desfontainea at first sight look rather different, they have a number of similarities, including some possible apomorphies, in common, and the circumscription adopted here seems reasonable.

Previous Relationships. In the first seven versions of this site (pre April 2008) Columelliaceae s.l. (= [Columelliaceae + Desfontainiaceae]) were placed immediately before the ordinal characterisation of Dipsacales - it seemed to make morphological sense and there was some other evidence for this position.

Includes Bruniaceae, Columelliaceae.

Synonymy: Columelliales Doweld, Desfontainiales Takhtajan

BRUNIACEAE Candolle, nom. cons.   Back to Bruniales

Bruniaceae

Ericoid shrubs, with lignotubers; myricetin +, iridoids?; stomata cyclocytic, orientation transverse; leaves spiral, small, close-set, lamina linear, margins entire, stipules +, often minute, colleter-like (0); leaf blade margins entire, apex mucronate; inflorescence racemose or spicate; flowers often small, (4-merous); (K connate), C often clawed (connate); A (adnate to C), anthers ± elongate; nectary +/0; G ± inferior, ?oblique, placentation apical-axile, style branched to the base, stigma capitate; ovules 2/carpel, micropyle long, integument 9-10 cells across, endothelium 0, hypostase +; fruit achenial or nut-like, single-seeded, K persistent; seed (arillate), coat ?; endosperm type?, haustoria?.

6/81[list]/75. South Africa, almost entirely the Cape Province, also KwaZulu-Natal (map: from Claßen-Bockhoff 2000). [Photo - Flowers.]

Age. Crown-group Bruniaceae are dated to 99.5-59.7 m.y.a. (Quint & Claßen-Bockhoff 2008).

There is a striking similarity between the flowers of Actinocalyx, from the Upper Cretaceous (Santonian/Campanian) ca 83.6 m.y.a. of Sweden, and those of Bruniaceae (Hall 1987).

1. Linconieae Quint & Claßen-Bockhoff

Stomata with cuticular rim; petals hard in texture; anthers sagittate, apically connate, with sterile tip; (G [3]); (fruits with two seeds); n = ?

1/3. The Cape.

[Audouineae + Brunieae]: (styles also connate); x = 11.

2. Audouineae Niedenzu

(Stomata with cuticular rim); (flowers single; inflorescence involucrate); anther thecae parallel, connate and attached to the connective their entire length; (pollen 4-5-colpate); (G [1, 3]); (ovules to 8/carpel); n = 11.

2/14. Southwestern part of the Western Cape.

3. Brunieae Quint & Claßssen-Bockhoff

(Lignotubers 0); inflorescence capitate, (involucrate), (with terminal flower and development centrifugal), (flowers single), etc.; anthers versatile, thecae (to rotund); (pollen to 10-colporate); (G [1]), (superior); ovules 1(-10)/carpel; (fruit with fleshy accrescent K), (dehiscent, also splitting adaxially); (seeds arillate); n = 22, 23.

3/64. Brunia (37). Esp. the Western Cape (Eastern Cape; KwaZulu-Natal).

Synonymy: Berzeliaceae Nakai

Evolution. Divergence & Distribution. The three main clades (tribes) had all diverged by 33.7-20.2 m.y.a., yet much of the diversification within them has occurred within the last 18-3 m.y. (Quint & Claßen-Bockhoff 2008).

Pollination Biology & Seed Dispersal. Myrmechory occurs in this clade (Lengyel et al. 2010).

Chemistry, Morphology, etc. Bruniaceae are poorly known. The plant is reported to be scented and the leaf apex has suberised cells produced by a localised cork cambium, hence the black tip (Carlquist 1990a). The corolla "tube" is at least sometimes formed by adnation of the filaments to adjacent free petals, and the petals are developmentally initally free (Quint & Claßen-Bockhoff 2006b). The androecium is often weakly monosymmetric, the abaxial pair of stamens being larger than the others. Gynoecial variation is considerable; in genera like Berzelia where there is only a single carpel (really pseudomonomery?), there is only a single locule, ovule and style. There are only one or two layers of parietal cells.

Some information is taken from from Saxton (1910: no endothelium shown) and Mauritzon (1939a), both embryology, Jay (1968a: chemistry), Hall (1988: pollen), Dahlgren and van Wyk (1988), Gregory (1998: anatomy), Endress and Stumpf (1991), Claßen-Bockhoff (2000: detailed discussion of inflorescences), and Quint and Claßen-Bockhoff (2006b: floral ontogeny); for wood anatomy, see Carlquist (1978b).

Phylogeny. The phylogenetic stucture [Linconieae [Audoinieae + Brunieae]] is strongly supported (Quint & Claßen-Bockhoff 2006a).

Classification. Claßssen-Bockhoff et al. (2011) propose a tribal classification based on the phylogeny by Quint and Claßssen-Bockhoff (2006a). For a somewhat dated monograph, see Pillans (1947).

Previous Relationships. Bruniaceae have often been linked with the South African Grubbiaceae, here in Cornales (see Hall 1987 for some references), but the similarities between the two probably reflect the fact that they are ericoid shrubs growing in similar habitats.

COLUMELLIACEAE D. Don, nom. cons.   Back to Bruniales

Cork cambium deep-seated; pericyclic fibres 0; petiole bundles arcuate; bud without scales; leaves opposite, petiole bases meeting, lamina margins toothed; inflorescence terminal, cymose; flowers medium-sized; anther connective well-developed, thecae attached their entire length; ovules many/carpel, endothelium poorly developed; K persistent in fruit; seeds many, anticlinal exotestal cells much thickened, with plasmodesmata.

2[list]/5. Costa Rica to Chile, montane in the tropical part of this area.

Age. These two genera diverged around 64 m.y.a. (Bremer et al. 2004).

Columellia

1. Columellia D. Don

Shrubs or trees; plant bitter tasting, iridoids 0; styloids +; central petiole bundle much the largest; cuticle waxes as tubules; lamina (with glands on lower surface), teeth glandular (0); flowers (4-8-merous); K ± valvate; A 2, latrorse, thecae semicircular, connective broad, expanded; tapetum amoeboid; G largely inferior, placentation intrusive parietal or lobed-axile, stigma with two broad lobes, almost cup-like; ovules ?lacking parietal tissue; fruit septicidal and part loculicidal, opening down the sides; exotestal cells elongated; endosperm type?; n = ?

1/4. S. Colombia to Bolivia, in the Andes (map: from Brizicky 1961). [Photo - Flower.]

2. Desfontainia Ruíz & Pavón

Desfontainia

Shrubs; route I secoiridoids +, tannin 0; scattered sclereids in the pericyclic position; lamina vernation conduplicate, teeth spiny, colleters +; inflorescence usu. single-flowered; K connate at base, C contorted; A adnate in throat, filaments stout; G [5(-7)], opposite petals, placentation axile but apically parietal, stigma only slightly expanded; ovule with integument 6-9 cells across; fruit a berry; exotestal cells polygonal, outer walls pectic, other walls, especially anticlinal, lignified; endosperm haustoria small; n = 7.

1/1: Desfontainia spinosa. Costa Rica to Chile. [Photo - Flower.]

Symonymy: Desfontainiaceae Pfeiffer

Evolution. Divergence & Distribution. Another old, species-poor but morphologically quite diverse group that is restricted to South America.

Chemistry, Morphology, etc. The intervascular pits of Desfontainia are scalariform or circular and bordered, not vestured. All the cells in even the young stem are slightly lignified. Myricetin?

The two stamens of Columellia are the adaxial pair and have latrorse, semicircular thecae attached their length to an expanded connective; the anthers are described as being extrorse by Backlund and Donoghue (1996). Although both Desfontainia and Viburnum have similar, smooth pollen orbicules, these are also quite widely distributed in Gentianales, at least (Vinckier & Smets 2002). Maldonado de Magnano (1986a) suggests that the ovules of Desfontainea are weakly crassinucellate, with a single layer of nucellar cells between the megaspore and the nucellar epithelium; an endothelium is at most poorly developed, and there is a large haustorial suspensor.

For Columellia information was taken from Hasselberg (1937), Stern et al. (1969: anatomy), Backlund and Donoghue (1996: general), and Gregory (1998: anatomy), for waxes, see Theisen and Barthlott (1994) and Fehrenbach and Barthlott (1988: cuticle platelets as ribbons and rodlets): Further data from: Zak & Jaramillo 3266. For Desfontainia information was taken from Backlund and Donoghue (1996) and Hasselberg (1937): Further data from: Qin 710 and Zarucchi et al. 5195.

Previous Relationships. The relationships of Columelliaceae have long been uncertain; the family was placed in Rosales by Cronquist (1981) and in Hydrangeales by Takhtajan (1997). Desfontainea has usually been included in the heterogeneous Loganiaceae - and so entirely unrelated.