EMBRYOPSIDA Pirani & Prado

Gametophyte dominant, independent, multicellular, thalloid, with single-celled apical meristem, showing gravitropism; rhizoids +, unicellular; flavonoids + [absorbtion of UV radiation]; chloroplasts lacking pyrenoids; protoplasm dessication tolerant [plant poikilohydric]; cuticle +; cell walls with (1->4)-ß-D-glucans [xyloglucans], lignin +; several chloroplasts per cell; 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, centrioles develop de novo, associated with basal bodies of flagellae, multilayered structure +, proximal end of basal bodies lacking symmetry, stellate pattern associated with doublet tubules of transition zone; spermatozoids with a left-handed coil; male gametes with 2 lateral flagellae; 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, wall with several trilamellar layers [white-line centred layers, i.e. walls multilamellate]; close association between the trnLUAA and trnFGAA genes on the chloroplast genome.

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 common ancestor of the group.

STOMATOPHYTES

Abscisic acid, ?D-methionine +; sporangium with seta, seta developing from basal meristem [between epibasal and hypobasal cells], sporangial columella + [developing from endothecial 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; 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; spore walls not multilamellate [?here].

EXTANT TRACHEOPHYTA / VASCULAR PLANTS

Photosynthetic red light response; water content of protoplasm relatively stable [plant homoiohydric]; control of leaf hydration passive; (condensed or nonhydrolyzable tannins/proanthocyanidins +); sporophyte soon independent, dominant, with basipetal polar auxin transport; vascular tissue +, sieve cells + [nucleus degenerating], tracheids +, in both protoxylem and metaxylem; endodermis +; root xylem exarch [development centripetal]; 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; stellate pattern split between doublet and triplet regions of transition zone; placenta with single layer of transfer cells in both sporophytic and gametophytic generations, embryonic axis not straight [root lateral with respect to the longitudinal axis; plant homorhizic].

[MONILOPHYTA + LIGNOPHYTA]

Branching ± indeterminate; lateral roots +, endogenous, root apex multicellular, root cap +; 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; male gametes multiflagellate, basal bodies staggered, blepharoplasts paired; chloroplast long single copy ca 30kb inversion [from psbM to ycf2].

LIGNOPHYTA†

Plant 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].

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 derived from (some) sinapyl and particularly coniferyl alcohols [hence with p-hydroxyphenyl and guaiacyl lignin units, so no Maüle reaction]; root stele with xylem and phloem originating on alternate radii, not medullated [no pith], cork cambium deep seated; arbuscular mycorrhizae +; 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 +; stem 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.; leaves with petiole and lamina, development basipetal, blade simple; branches axillary (buds not associated with all leaves), exogenous; prophylls two, lateral; plant heterosporous, sporangia borne on sporophylls; microsporophylls aggregated in indeterminate cones/strobili; true pollen +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, parietal tissue 2+ cells across, megaspore tetrad linear, functional megaspore single, chalazal, lacking sporopollenin, megasporangium indehiscent; pollen grains landing on ovule; gametophytes dependent on sporophyte; male gametophyte development initially endosporic, tube developing from distal end of grain, gametes two, developing after pollination, with cell walls; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large" [ca 8 mm3], but not much bigger than ovule, with morphological dormancy; 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], white, 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?], S [syringyl] lignin units common [positive Maüle reaction - syringyl:guaiacyl ratio more than 2-2.5:1], and 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, 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, 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], ± embedded in the filament, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally, 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 +, thin, compact, lamellate only in the apertural regions; nectary 0; carpels present, superior, free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, dry [not secretory]; 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]; supra-stylar extra-gynoecial compitum +; ovule not increasing in size between pollination and fertilization; pollen grains landing on stigma, bicellular at dispersal, mature male gametophyte tricellular, germinating in less than 3 hours, pollination siphonogamous, tube elongated, 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, flagellae 0, double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; seed exotestal, much larger than ovule at time of fertilization; 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]; 2C genome size 1-8.2 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; 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; 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]]]]: ?

Age. The appproximate age for this node is (106-)96(-85) m.y. (Beaulieu et al. 2013a). Bell et al. (2010: note position of Bruniales) suggested an age (96-)84, 74(-65) m.y., other ages are suggested in the context of topolgies different from that followed here.

Divergence & Distribution. Initial diversification probably occurred in the southern hemisphere (Beaulieu et al. 2013a).

Phylogeny. Various small families have been placed around here in the campanulids, but initially with uncertain support; recent work is clarifying their relationships (Winkworth et al. 2008a; especially Tank & Donoghue 2010; Beaulieu et al. 2013b). For Polyosmaceae and Escalloniaceae, see below. Paracryphiaceae form a clade sister to Dipsacales, thus the clade [Paracryphia + Quintinia] was sister to Dipsacales, although not in analyses that included coding chloroplast genes (Winkworth et al. 2008a).

In earlier versions of this site (pre April 2008), [Columelliaceae + Desfontainiaceae] (= Columelliaceae s.l.) were placed sister to Dipsacales, with which they show some morphological similarities. The position of Columelliaceae s.l. in that area had been suggested by Bremer et al. (2001) and especially by Lundberg (2001e; see also Backlund 1996), although support was at best moderate. Both Columelliaceae s.l. and Dipsacales have opposite leaves, and Columellia has amoeboid tapetum (c.f. Bremer et al. 2001) like Dipsacales although Desfontainia does not (Maldonado de Magnano 1986a). The similarities that the pair has with Dipsacales may indicate either substantial homoplasy or (less likely) a suite of rather basal synapomorphies in the campanulids of which there is currently no indication. Other relationships have been suggested (Gustafsson et al. 1996; Backlund & Bremer 1997; Pyck & Smets 2000; Bell et al. 2001); for the possible association of Bruniaceae with Asterales, see Lundberg (2001e). Here Columelliaceae s.l. together with Bruniaceae make up Bruniales.

For relationships of the larger families, see the asterid IIclade. The overall topology [Bruniales [Apiales [Paracryphiales + Dipsacales]]] seem to be fairly well established (Winkworth et al. 2008a; Tank & Donoghue 2010; Beaulieu et al. 2013a); however, the position of Escalloniales is sometimes rather poorly supported.

ESCALLONIALES Martius  Main Tree.

Inflorescence racemose; petals free; A adnate or not to C, anthers basifixed; nectary +; ovule with integument 5-10 cells across. - 1 family, 9 genera, 130 species.

Note: 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. For relationships in the order, see below. In a 17-gene analysis, Polyosma linked with Quintinia, in Paracryphiales here (see Soltis et al. 2011, and q.v. for a possible explanation).

Includes Escalloniaceae.

Synonymy: Escalloniineae Shipunov - Tribelales Doweld

ESCALLONIACEAE Dumortier, nom. cons.   Back to Escalloniales

Plants Al accumulators; inflorescence racemose; ovary inferior; parietal tissue ca 1 cell across[?].

9[list] /130. Réunion, E. Himalayas and S. China to E. Australia and New Zealand, Central and South America.

Age. The crown age for this family is in excess of 110 m.y.a. (Bremer et al. 2004: note topology).

Polyosma

1. Polyosma Blume

Trees; iridoids +; wood storying +; pericycle with sclereids; nodes 1:1; petiole bundle?; stomata ?; hairs unicellular; leaves opposite, lamina margins toothed or not; flowers 4-merous; K connate, C valvate; pollen triporate; G placentation intrusive parietal, stigma capitate, bilobed; ovules many/carpel, straight, integument 4-10 cells across, ?endothelium, parietal tissue ca 1 cell across[?], nucellus base massive; (megaspore mother cells several); fruit a 1-seeded drupe; endosperm starchy, thick-walled, ?haustoria, embryo undifferentiated; n = ?

1/60. E. Himalayas and S. China to E. Australia and New Caledonia (map: in part from GBIF Data Portal 3.i.2009).

Synoymy: Polyosmaceae Blume

2. Escallonia, etc.

Escalloniaceae

Shrubs to trees (annual herbs); route I seco- and route II decarboxylated iridoids, flavonols +; (cork cambium deep-seated - Escallonia); nodes 3:3 (1:1, 5:5); petiole bundle arcuate; heads of glandular hairs with radially arranged cells; stomata anomocytic; leaves spiral, lamina vernation supervolute (conduplicate), margins with broad glandular teeth and two accessory veins (entire); (inflorescence a cyme - Eremosyne; flowers single, terminal - Tribeles); flowers 5-9-merous; C contorted, (± connate - Escallonia); anthers longer than connective (short - Tribeles), placentoid +; ?pollen; G [2-4], (largely superior), collateral, or median member adaxial, placentation also basal, parietal, style ± branched or not, stigma punctate to capitate or clavate-lobed, wet; ovule 1-many/carpel, (ascending), integument 5-8 cells across, micropyle long, (ovule bitegmic - Tribeles), endothelium +/weak; embryo sac protruding at micropyle; fruit a septicidal capsule, often splitting down the sides; exotestal cells with inner walls thickened and lignified (not), elongated or not (palisade - Tribeles), meso- and/or endotesta usu. persist; micropylar endosperm haustoria +, (embryo long); n = 12.

8/68: Escallonia (40). Scattered, but largely southern: Central and South America, SE and SW Australia (Eremosyne pectinata), New Zealand, Réunion (map: from Sleumer 1968; FloraBase 2007). [Photos - Escallonia Flower, Valdivia Flower.]

Age. The appproximate crown-group age for this clade is (85-)80, 72(-67) m.y. - see Wikström et al. (2001), or (87-)72, 65(-48) m.y. - see Bell et al. (2010). Since in both cases the age is of the [Eremosyne + Escallonia] node (see below), this would be an underestimate of the age even of the immediate clade above.

Synonymy: Anopteraceae Doweld, Eremosynaceae Dandy, Tribelaceae Airy Shaw

Chemistry, Morphology, etc. This is a very heterogeneous and poorly known group; there is no information on endosperm development, etc. As Bensel and Palser (1975d, p. 693, see also 1975c) noted of Escallonioideae (for them, also including Quintinia [= Paracryphiales-Paracryphiaceae]), "The only conclusion that can be drawn about Escallonioideae is that it has been too little investigated in all aspects" - a conclusion that has so far unfortunately stood the test of time.

Forgesia has sclereids alone in the pericyclic sheath. Escallonia occasionally has prickles in the stipular position. Anopterus has parietal placentation, apparently a long micropyle, and winged seeds; the exotestal cells are elongated, and have thickened inner walls. Escallonia lacks the mitochondrial coxII.i3 intron; other taxa have not been sampled.

For general anatomy, see Gornall et al. (1998), for some information on ovules, see Mauritzon (1933). For anatomy of Forgesia, see Ramamonjiarisoa (1980: young stem with a complete vascular cylinder). For Escallonia, see Stern (1974: anatomy), Swamy (1954: nodes), Al-Shammary and Gornal (1994: indumentum), and Krach (1976) and Nemirovich-Danchenko and Lobova (1998), both seeds, and Schnizlein (1843-1870: fam. 170: floral orientation). Much general information is taken from Lundberg (2001c: Polyosmaceae, 2001d: Escalloniaceae), while for some information on embryology and seed, see Mauritzon (1933, 1938a) and Danilova (1996), and for a summary of information on Eremosyne (?ovules not quite tenuinucellate), see Hibsch-Jetter et al. (1997).

Phylogeny. A relationship between Escalloniaceae s. str. and Eremosynaceae has strong 3-gene support (Soltis et al. 2000; see also Hibsch-Jetter et al. 1997); a further association with Tribelaceae is only weakly supported (Savolainen et al. 2000b: rbcL alone). However, Escalloniaceae were found to be paraphyletic if Eremosynaceae and Tribelaceae were excluded (or Anopterus would have to be in a separate family), furthermore, the monogeneric Polyosmaceae were sister to the whole group (Lundberg 2001e: three-gene Bayesian analysis); Bremer et al. (2004) recovered the relationships [[Tribeles + Polyosma] [Escallonia + Eremosyne]]. Details of relationships between the genera remain unclear (Tank & Donoghue 2010, see also Beaulieu et al. 2013a, but sampling). Sede et al. (2013) found strong support for [Valdivia (Chile) + Forgesia (Reunion)] as sister to Escallonia (see also Tank & Donoghue 2010; Zapata 2012); although Eremosyne was sister to the rest of the family, apparent differences from Bremer et al. (2004) may be a rooting problem.

For the phylogeny of Escallonia, see Zapata (2013) and Sede et al. (2013), there is considerable geographical structuring of relationships.

Classification. Lundberg (pers. comm.) thought that all these families should be combined into one, and so they are (see also A.P.G. III 2009).

Previous Relationships. Krach (1976, 1977) suggested that Escalloniaceae, in which he included Abrophyllaceae and Argophyllum (Rousseaceae), both in Asterales here, were close to Hydrangeaceae, and should be placed in an Escalloniales; Escalloniaceae were placed in Hydrangeales by Takhtajan 1997), although he wasn't sure about the position of Anopterus. Eremosyne was previously frequently included in Saxifragaceae (e.g. Cronquist 1981) or Saxifragales (Takhtajan 1997). Tribeles was placed in Hydrangeales by Takhtajan (1997), who described the capsule as being loculicidal.

Thanks. I am grateful to F. Zapata for discussion.