EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline; primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem complex; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5(-8) mm/mm²; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication, mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, cyanogenesis via tyrosine pathway [ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less 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; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with sieve plate, companion cells from same mother cell that gave rise to the tube, the sieve tube with P-proteins; nodes unilacunar; stomata with ends of guard cells level with aperture, paracytic; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, vein endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable, P not differentiated, outer members not enclosing the rest of the bud, smaller than inner members, A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther, tapetum glandular, binucleate, microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing, pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, exine columellar, endexine thin, compact, lamellate only in the apertural regions, pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm, nectary 0, G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte ?type, stylulus short, hollow, stigma ± decurrent, wet [secretory]; P deciduous in fruit; seed exotestal; double fertilisation +, endosperm ?diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, 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 PHYA/PHYC gene pairs.

Possible apomorphies are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied. Furthermore, details of relationships among gymnosperms will affect the level at which some of these characters are pegged.

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates; nucleus of egg cell sister to one of the polar nuclei; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

MALPIGHIALES Martius  Main Tree, Synapomorphies.

Vessel element type?; (sieve tubes with non-dispersive protein bodies); leaf margin toothed [teeth with a single vein running into a congested ± deciduous apex], stipules +; stigma dry; exotegmen fibrous. - 39 families, 716 genera, 15935 species.

Crown Malpighiales probably diverged some time in the Cretaceous-late Aptian, perhaps some 114 million years before present ([119.4-]113.8[-110.7]/[105.9-]101.6[-101.1] million years before present - high and low estimates: Davis et al. 2005a); diversification seems to have been rapid (Wikström et al. 2001, however, suggest divergence dates of 91-88 million years before present, diversification beginning only 81-77 million years before present, with stem groups of many families being evident before the beginning of the Tertiary). The order contains ca 7.8% eudicot diversity (Magallón et al. 1999) and is particularly important in tropical rainforests where it is an important component of the diversity of the understory; it accounts for up to some 28% of the species and 38% of the total stems (Davis et al. 2005a); members of Ericales are the other main component.

Caterpillars of outgroups to Nymphalidae-Nymphalinae, -Melitaeini, etc., are quite common on Mapighiales (Nylin & Wahlberg 2008). The butterfly Cymothoë has hosts widely scattered in this order (Ackery 1988), although also found on Bignoniaceae (one species) and Rhamnaceae (sometimes another species). Phyllonorycter leaf-mining moths (Lepidoptera-Gracillariidae - Phyllocnistinae) seem to have diversified on this clade (and especially Fagales) some time in the region of 50.8-27.3 million years before present, well after the Malpighiales diversified, and after the genus itself evolved, some 76.3-50.3 million years before present (Lopez-Vaamonde et al. 2006).

Articulation of the pedicels is another feature that may be common to the order. The atpF gene has been lost several times in this clade, alone among angiosperms, however, details of its distribution remain unclear due to poor sampling and lack of phylogenetic relaolution in the clade (Daniell et al. 2008).

Although Malpighiales are strongly supported as being monophyletic (e.g. Davis et al. 2005a), relationships within them are still poorly understood (e.g. Soltis et al. 2007a). Those in the tree shown here are mostly taken from Litt and Chase (1999), Schwarzbach and Ricklefs (2000), Chase et al. (2002), and Davis and Chase (2004), but they are in general agreement with relationships apparent in broader studies. Davis et al. (2005a) have recently clarified some relationships in Malpighiales in a four-gene (all three compartments) analysis, in particular suggesting an association between the families with parietal placentation (and also Goupiaceae) and that Centroplacus should be recognised as a separate family (see also Korotkova et al. 2007 for relationships in Malpighiales). The inclusion of Rafflesiaceae in Malpighiales follows the recent findings of Barkman et al. (2004, 2007), Davis and Wurdack (2004), and in particular Davis et al. (2006), who place it with strong support as sister to Euphorbiaceae s. str. It seems useful to adopt a narrow circumscription for families that used to be included in Flacourtiaceae and Euphorbiaceae s.l. Even if future work suggests reaggregation of genera that used to be placed in these two families, groupings within these new units will be different from those suggested by previous classifications. Note that the realignments caused by the break-up of the old Flacourtiaceae and integration with Salicaceae and Achariaceae correlate well with a number of morphological/anatomical characters. Paracytic stomata may characterise a sizeable clade in Malpighiales, and three-carpellate gynoecia are common...

See Endress and Matthews (2006b) for petal appendages, etc., in the order, also Matthews and Endress (2008) discuss other floral variation; Tokuoka and Tobe (2006) integrate testa anatomy and embryology with phylogeny.

Includes Achariaceae, Balanopaceae, Bhesa, Bonnetiaceae, Caryocaraceae, Centroplacaceae, Chrysobalanaceae, Clusiaceae, Ctenolophonaceae, Dichapetalaceae, Elatinaceae, Erythroxylaceae, Euphorbiaceae, Euphroniaceae, Goupiaceae, Humiriaceae, Hypericaceae, Irvingiaceae, Ixonanthaceae, Lacistemataceae, Linaceae, Lophopyxidaceae, Malpighiaceae, Malesherbiaceae, Medusagynaceae, Ochnaceae, Pandaceae, Passifloraceae, Peraceae, Phyllanthaceae, Picrodendraceae, Podostemaceae, Putranjivaceae, Quiinaceae, Rafflesiaceae, Rhizophoraceae, Salicaceae, Trigoniaceae, Turneraceae, Violaceae.

Synonymy: Balanopales Engler, Chrysobalanales Reveal & Doweld, Elatinales Nakai, Euphorbiales Lindley, Flacourtiales Heinze, Homaliales Bromhead, Huales Doweld, Hypericales Dumortier, Irvingiales Doweld, Lacistematales Baskerville, Linales Baskerville, Rafflesiales Oliver, Marathrales Dumortier, Medusagynales Reveal & Doweld, Ochnales Reveal, Pandales Engler & Gilg, Passiflorales Dumortier, Phyllanthales Doweld, Podostemales Lindley, Rhizophorales (Persoon) Reveal & Doweld, Salicales Lindley, Samydales Dumortier, Scyphostegiales Croizat, Turnerales Dumortier, Violales Perleb - Euphorbianae Reveal, Ochnanae Doweld, Podostemanae Reveal, Rafflesianae Reveal, Rhizophoranae Reveal & Doweld, Violanae Reveal - Malpighiopsida Bartling, Passifloropsida Brongniart, Podostemopsida G. Cusset & C. Cusset, Salicopsida Bartling, Violopsida Brongniart

[Achariaceae [Goupiaceae [Violaceae [Malesherbiaceae [Turneraceae + Passifloraceae]]] [Lacistemataceae + Salicaceae]]]: sieve tubes with non-dispersive protein bodies; cuticle waxes usu. 0; (foliar glands +); pedicels articulated; nectariferous tissue, stamens = and opposite sepals, G with median member abaxial, placentation parietal; seed arillate; endotegmen persistent; endosperm persistent, oily.

All six major clades in this group (Malesherbiaceae + Turneraceae + Passifloraceae as a single clade) may have diverged in the Cretaceous-Albian 111-100 million years before present, or somewhat later (Davis et al. 2005a, details are given for the individual clades: Wikström et al. 2001 suggests many of the clades do not diverge until [well] after 63 million years before present). Larvae of butterflies such as Nymphalidae-Acraeinae and N.-Nymphalinae-Heliconiini, -Vagrantini and -Argynnini commonly eat members of this group (Ehrlich & Raven 1964; see also Dahlgren & van Wyk 1988; Arbo 2006; Simonsen 2006; Silva-Brandão et al. 2008; Nylin & Wahlberg 2008; etc.); this is also discussed under individual families below. Some Acraeinae in particular may cue on the presence of the cyanogenic glucoside gynocardin in potential food plants, indeed, that larvae of Acraea horta, normally living on the woody Kiggelaria africana, ate herbaceous Achariaceae prompted the successfull search for that compound in the latter family (Steyn et al. 2002). Toxic compounds like gynocardin may be sequestered by the larva and passed on to the adult.

There are numerous anatomical, chemical and floral links between Salicaceae, Achariaceae and Violaceae, sometimes also Passifloraceae, Malesherbiaceae and Turneraceae (Nandi et al. 1998). Thus Achariaceae + Malesherbiaceae + Turneraceae + Passifloraceae have in common cyclopentenoid cyanogenic glucosides and/or cyclopentenyl fatty acids and also commonly some sort of corona or scales on petals. Molecular evidence that the whole group formed by families with parietal placentation is monophyletic has previously not been compelling (e.g. see Savolainen et al. 2000a; Chase et al. 2002), although part of the rpS 16 gene is absent from Passifloraceae, Violaceae, Salicaceae s. str., and Turneraceae (and also Linaceae and Malpighiaceae, so really a feature of Malpighiales?: see Downie & Palmer 1992). Salicaceae are weakly associated with Passifloraceae, and in turn with Humiriaceae and Pandaceae, in Chase et al (2002). Associations like Salicaceae + Passsifloraceae s.l. (inc. Turneraceae and Malesherbiaceae) are weakly supported, as are Achariaceae + Violaceae (and Lacistemataceae + Ctenolophonaceae)- see below. Tokuoka and Tobe (2006) found a weakly-supported relationship between the Passifloraceae group and Violaceae (see also Soltis et al. 2007a), and strongly supported relationships between Lacistemataceae and Salicaceae. However, Davis et al. (2005a) found a moderately supported association of these taxa with parietal placentation (59% bootstrap, 100% posterior probability), and also Goupiaceae, with axile placentation, and a similar grouping is also evident in e.g. Wikström et al. (2001) and Korotkova et al. (2007: 83% jacknife, 1.00 pp, Goupiaceae not included). Ixonanthes was rather surprisingly embedded in Achariaceae in the Bayesian analysis of Soltis et al. (2007a), but that was due to a misidentification of the material examined, which was a species of Hydnocarpus (K. Wurdack, pers. comm.).

There is much information on seed anatomy in Takhtajan (1992) while Krosnick et al. (2006) briefly discuss the evolution of polyandry in this group - in some cases, at least, the numerous stamens form a single whorl.

ACHARIACEAE Harms, nom. cons.   Back to Malpighiales

Shrubs to trees ((climbing) herbs); cyclopentenoid cyanogenic glucosides and/or cyclopentenyl fatty acids +, gynocardin, ellagic acid [Kiggelaria] +; vessel elements with simple or scalariform perforation plates; fibers septate; petiole bundle annular, with two lateral/adaxial strands (inverted medullary plate - Lindackeria); leaves spiral or two-ranked, margins entire (serrate; stipules 0), petiole often geniculate; (plant dioecious); inflorescence spicate or cymose (fasciculate); K and C spiral or not, not in a simple alternating relationship, K 2-5, C 4-15 (3-4, connate - Acharieae), often in two series, (adaxial scales +), A 5-many, opposite petals or irregular, centripetal or simultaneous, (from a ring meristem), anthers basifixed, elongate (barely so - Chiangodendron; porose; locellate), pollen also tricolporoidate, disc 0, G [2-10], median member?, ovules sessile, (atropous - Xylotheca, Hydnocarpus, Lindackeria; 1/carpel), (archesporium bicellular - Caloncoba) (embryo sac Allium-type [bisporic, 8-nucleate] - Acharieae), integuments 5-6 cell layers across, micropyle endo- or bistomal or zigzag, nucellar cap and epistase +, style (short), branched or not, stigma capitate-peltate; fruit also a berry; seed (arillate), coat thick, pachychalazal [not Hydnocarpus laurifolius], or both integuments multiplicative [possibly simply different interpretations], testa vascularised, sarcotestal and with stomata (Acharieae) or not, (inner mesotesta sclereidal), endotesta lignified, cells sclereidal (radially elongated); endosperm copious, suspensor 0, embryo green; n = 10, 12, 23.

30[list]/145: Hydnocarpus (40). Pantropical. (Map: from Sleumer 1954, 1980; Şerban Procheş, pers. comm. [Africa]; Andrew Ford, pers. comm. [Australia])[Photo - Flower, Fruit, Fruit, Acharia tragodes - Leaves.]

• Achariaceae can be recognised by their stipulate leaves, often entire leaf blades and pulvinate petioles, and flowers in which the number of sepals and petals is often clearly not the same, and/or the perianth is not simply biseriate, and/or the petals have basal, adaxial scales. There is no nectary, and the anthers are usually long. The fruits are often quite large, the placentation is parietal, and the seeds often have conspicuous vascular bundles in the coat.

The feeding behaviour of Acraeini butterfly larvae are consistent with the expanded family limits adopted here (Steyn et al. 2002, 2003 and references).

Circumscription (see Chase et al. 2002; Sosa et al. 2003): Acharieae (more or less herbaceous and viny; no testal bundles; zig-zag micropyle; fibrous exotegmen), Erythrospermeae (Erythrospermum - fibrous exotegmen), Pangieae (inc. Kiggelarieae) and Lindackerieae (Oncobeae minus Oncoba). Note that the family is divided into three strongly-supported clades, largely Hydnocarpus, Erythrospermeae + Lindackerieae, and Acharieae + Pangieae, and support for monophyly of the family as a whole is strong (Sosa et al. 2003).

Gynocardin is found in some other families of this group (Webber & Miller 2008). There are large and medium intervascular pits; the wood also has solitary pores and lacks tracheids. Lindackeria has superficial cork cambium. Pollen variation is considerable (Wendt 1988).

The bulk of Achariaceae have almost universally been included in Flacourtiaceae s.l. (Cronquist 1981; Takhtajan 1997) = Salicaceae, q.v. for relationships.

Information is taken from Hegnauer (1966, 1989, as Flacourtiaceae: chemistry), van Heel (1977, 1979: testa anatomy), Endress and Voser (1975: floral development), Miller (1975: wood anatomy), Spencer and Seigler (1985: chemistry), Lemke (1988), Gavrilova (1998: pollen) and Steyn et al. (2002a, b, 2003: ovule development, testa anatomy); see also Judd (1997a) and especially Chase et al. (2002). Bernhard and Endress (1999) discuss androecial initiation. I thank Sue Zmarzty for comments.

Synonymy: Erythrospermaceae Doweld, Kiggelariaceae Link, Pangiaceae Endlicher

VIOLACEAE Batsch, nom. cons.   Back to Malpighiales

Vessel elements with simple or scalariform perforation plates; calcium oxalate often as crystals; petiole bundles arcuate; stomata para- or anisocytic; pedicels articulated; K quincuncial, A with abaxial nectary, G (3]; K persistent in fruit; exotesta subpalisade to tabular, ± thickened, (mesotesta sclerenchymatous), endotesta usu. crystalliferous.

23[list]/800. World-wide (Map: from Hultén 1958, 1971; George 1982; Hultén & Fries 1986; Hekking 1988 - incomplete for South America).

1. Fusipermoideae Hekking

C contorted, fleshy annular 5-lobed disc, filaments adnate to inner surface at indentations, A with minute fringed apical scales, thecae cordate/trapezoid, confluent apically?; capsule ca 2 mm long; seeds arillate/carunculate.

1/3. Panama, Columbia, Peru.

For information, see Cuatrecasas (1950) and Hekking (1984), the former describes the scales as being ventral appendages of the connective. Whether or not there are really two scales is unclear.

2. Violoideae

Herbs to trees (lianes); plants often Al accumulators; tannins 0 [woody members?]; vessel elements with simple or scalariform perforation plates; calcium oxalate often as crystals; petiole bundles arcuate; stomata para- or anisocytic; leaves spiral or two-ranked (opposite), margins involute, colleters +, (stipules petiolar; lobed); flowers poly- or monosymmetric, C quincuncial, abaxial C spurred or not, A (3), all or 2 abaxial nectariferous, (filaments connate), anthers connivent, (thecae horizontal), connectives prolonged (± 0), G [(2-5)], micropyle zigzag (endostomal), hypostase +, styles separate or style +, stigmatic head subcapitate, asymmetrical or not, receptive area small; (fruit a berry, nutlike); seeds (winged) often arillate/carunculate, exotesta subpalisade to tabular, ± thickened, (mesotesta sclerenchymatous), endotesta usu. crystalliferous; embryo (small), green [Viola]; n = 6-13+.

22/795: Viola (400-600: cleistogamy widespread, V. tricolor, the pansy, and the related V. arvensis important in early studies of genetics and speciation), Rinorea (160-270), Hybanthus (90-150). World-wide; woody taxa esp. in the lowland tropics. [Photo - Leonia, Alexis fruit and flowers, Viola.]

• Violaceae may be recognised by their serrate, stipulate leaves which often dry yellowish and with raised reticulum, their flowers, which are often spurred and have stamens with short filaments and very well-developed connectives that closely surround the gynoecium, and their fruit, which is often a distinctive loculicidal capsule, the rather large seeds being dispersed explosively as the fruit wall closes round them and squeezes them out. Plants are herbs, trees or lianes.

Violaceae are the preferred hosts of the majority of fritillaries (Nymphalidae: Argynnini - see Simonsen 2006).

Melicytus is woody, dioecious, and baccate; the flowers are almost radially symmetrical... Not surprisingly, woody Violaceae are quite commonly often wrongly identified or not named at all, as Ron Leisner will confirm - however, the vegetative characters mentioned above do help. Viola has storied cambium. In Anchietea and Decorsella the seeds mature exposed on the open carpels.

Violaceae are weakly associated with Achariaceae (and Goupiaceae, Lacistemataceae and Ctenolophonaceae) in Chase et al (2002a).

There is good support for the relationships [Fusispermum [Rinorea + the rest]], with Leonioideae being embedded within Violoideae (Tokuoka 2008, see also Feng & Ballard 2005).

For embryology, etc., see Singh (1970), for chemistry, see Hegnauer (1970, 1990), for general information, see Hekking (1988) and Munzinger and Ballard (2003: also key to genera, two undescribed).

Synonymy: Alsodeiaceae J. Agardh, Leoniaceae A. L. de Candolle

SALICACEAE Mirbel, nom. cons.   Back to Malpighiales

(Ectomycorrhizal) evergreen (deciduous) trees; cocarcinogen, (gynocardin, ellagic acid +), tanniniferous; cork?; vessel elements with simple or scalariform perforation plates; (nodes 2:2 - some Azara); petiole bundle arcuate or annular with flange bundles; stomata ?; leaves spiral or two-ranked (opposite), supervolute-curved or involute, teeth (margin entire; venation palmate; glands +; stipules 0); (plant dioecious), inflorescence various; flowers often small, 3-6-merous, (hypanthium +), K (0-)3-8(-15), often valvate (connate), C 0 or = K (more than K - e.g. some Scolopieae; basally connate; corona +), disc often with glands or lobes (the latter intrastaminal; 0), A 1 to many (fasciculate, opposite petals), centrifugal (simultaneous) initiation, anthers (extrorse), ellipsoid to subglobose (linear), (pollen inaperturate - Populus), G [2-5(-13)], (to inferior), (placentation axile; basal), ovules (unitegmic) straight [atropous] (anatropous), (nucellar cap +), (outer and inner integuments ca 5 cells across), micropyle (exo - Idesia)/bistomal/zig-zag, embryo sac elongated, ± protruding into the micropyle, (bisporic, Allium-type), styles separate or style +; fruit also a berry (drupe); seeds arillate (with hairs [of arillate origin], testa also multiplicative but otherwise undistinguished, or exotesta or endotesta [Oncoba] palisade); endosperm + (0 - Salix), (embryo green); n = 9, 10-12, 19.

55[list]/1010: Salix (450: notorious for interspecific hybridisation; connate P modified as nectary, vascular traces to absent members, secondarily insect-pollinated?), Casearia (180), Homalium (180), Xylosma (85). Pantropical, also temperate (but few Australia, not New Zealand) to Arctic (Map: (Africa incomplete) from Sleumer 1954; Meusel et al. 1975; Sleumer 1980; Hultén & Fries 1986). [Photo - Flower, Fruit.]

• Salicaceae can be recognised by their alternate, serrate, stipulate leaves with distinctive "salicoid teeth"; these latter have a small vein proceeding into the tooth, where it expands, the apex of the tooth being a variously coloured spherical gland or stout hair. The flowers often have many spreading stamens and a commonly three-carpellate gynoecium with parietal placentation and either separate styles or a long, persistent style; the fruit is often a capsule. When there are both sepals and petals, the two whorls are in a regular alternating arrangement, and the nectariferous disc or nectar glands and often outside the stamens. A number of taxa have pellucid foliar glands (Abatia, Casearia, Ryania, Zuelania).

Ehrlich and Raven (1964) noted that Atella (Nymphalinae) feeds on Flacourtiaceae and Salicaceae, while some Notodontidae moths (Miller 1992), rusts, e.g. Melampsora spp. on Salix, M. idesiae on Idesia (Holm 1979), etc., show similar host pattern. Boucher et al. (2003) describe Pseudosalix, an Eocene fossil from North America, which is morphologically intermediate between Salix and more morphologically conventional Salicaceae.

Circumscription: Abatieae, Bembicieae, Prockieae (inc. Banareae), Oncobeae (Oncoba only), Homalieae, Saliceae, Samydeae (Casearieae), Scolopieae, and Scyphostegieae (see Chase et al. 2002). Tribal limits may well have to be adjusted, thus Saliceae will probably have to be expanded, Flacourtieae in their current circumscription are polyphlyetic, etc. Trichostephanus (Trichostephaneae) was not assigned to any family (Chase et al. 2002), but in its absence of petals and in having a disc at the base of the calyx it is like Casearia. Oncoba is remarkably like other members of the erstwhile Oncobeae (see now Achariaceae - Lindackerieae), but they differ in chemistry, leaf tooth type, and stamen initiation.

Alford (2003) recognised three families for the New World flacs in addition to Achariaceae, Berberidopsidaceae and Lacistemataceae. His Samydaceae have arillate seeds, punctate or lineate leaves, and flowers with a hypanthium, while the two other families have salicoid leaf teeth, but lack arillate seeds or hypanthial flowers. Salicaeae have dilated stigmas and petals and sometimes sepals absent, while Flacourtiaceae s. str. have attenuate, lobed, or capitate stigmas and petals present or absent.

"Flacourtiaceae as a family is only a fiction; only the tribes are homogeneous" (Sleumer, in Miller 1975). Salicaceae s. str. (cuticle waxes as platelet rosettes, or 0; inflorescences catkins; P much reduced, G [2], collateral, ovules usu. unitegmic; seeds with long placental hairs at base, tegmen not persisting; embryo green) seem quite distinctive. There is a gene duplication in the common ancestor of Salix and Populus, the salicoid duplication, dated to 65-60 million years before present (Tuskan et al. 2006); it will be interesting to know if other Salicaceae s.l. have this duplication. Indeed, it has been observed that Salicaceae s. str. and Flacourtiaceae-Idesieae are very close - they have distinctive leaf teeth, phenolic-type compounds such as salicin are found here only, etc., and as just noted, rusts and caterpillars, perhaps keying in on chemical characters, show similar distributions (see e.g. Meeuse 1975). Elongated embryo sacs occur in both Salicaceae and old Flacourtiaceae (Steyn et al. 2005). Furthermore, it was known that within the old Flacourtiaceae there were two rather different kinds of seed coat (Corner 1976). Chase et al. (2002) have clarified the situation (see also Judd 1997a; Nandi et al. 1998; T. Azuma et al. 2000; Savolainen et al. 2000a), although sampling within tribes still needs to be extended. Some of the old Flacourtiaceae are now in Achariaceae, a few in Lacistemataceae, while Berberidopsideae are in Berberidopsidales (as Berberidopsidaceae) and Aphloia (Aphloiaceae) is placed in Crossosomatales. The remainder of Flacourtiaceae are here. Variation in chemistry, leaf teeth, floral morphology, and seed coat anatomy is largely correlated with this division. Salicoid teeth are quite variable, but all have palisade cells over parenchyma and are well supplied by vascular, especially xylem, tissue (Wilkinson 2007). It should also be noted that Casearia, which may lack salicoid leaf teeth and has apetalous flowers with the disc on the basal-adaxial surface of the calyx, is sister to the rest of Salicaceae, although support for this position is weak (Chase et al. 2002, but cf. D. Soltis et al. 1999, 2000).

Salix and its immediate relatives have ectomycorrhizae. Banara is the only genus reported to have cyanogenic glycosides, but it is well embedded within Salicaceae (Chase et al. 2002). The perforation plates of the tracheary elements are more or less simple and the intervascular pits are small. Xylosma and some Casearia seem to have unilacunar nodes. Leaf traces arise an internode below the leaf they innervate in Hasseltia. Casearia can have phyllanthoid branching, the orthotropic axes having spirally arranged and reduced leaves while the plagiotropic branches are sylleptic and have fully-expanded and two-ranked leaves. Xylosma, Flacourtia, etc. have groups of large sclereids in the phloem (Zahur 1959). Abatia has opposite leaves with at most very small stipules and marginal glands at the base of the lamina, its valvate perianth members are basally connate and bear many filamentous processes, and it lacks any nectary. There are also taxa with pli-nerved leaf blades and foliar glands, while Populus is dioecious and wind-pollinated... The disc is very variable, and it is often broken up into lobes; it is then sometimes intrastaminal. The embryo sac more or less protrudes into the micropyle in Archevaletaia (Mahshwari 1950). The exotegmen of Dovyalis consists of ribbon-type cells.

Some information is taken from Hegnauer (1973, 1990, also 1966, 1989, as Flacourtiaceae: chemistry), van Heel (1977, 1979: testa anatomy), Miller (1975: wood anatomy), Spencer and Seigler (1985: chemistry), Lemke (1988: general), Gavrilova (1998: pollen), Leskinen and Alström-Rapaport (1999: relationships of Salix and its immediate relatives), and Steyn et al. (2004, 2005: ovule and seed development); see also Judd (1997a) and especially Chase et al. (2002). Bernhard and Endress (1999) discuss androecial initiation. I am grateful to S. Zmarzty for comments.

Synonymy: Bembiciaceae R. C. Keating & Takhtajan, Caseariaceae Doweld, Flacourtiaceae Richard, Homaliaceae R. Brown (G inferior), Poliothyrsidaceae Doweld, Prockiaceae Bertuch, Samydaceae Ventenat, nom. cons., Scyphostegiaceae Hutchinson, nom. cons. (Cork superficial; vessels in radial multiples, with simple (and scalariform) perforations; rays mostly uniseriate; petiole bundle annular and with adaxial mass of xylem and phloem becoming adaxial inverted plate of vascular tissue; stomata paracytic; leaves distichous; plant dioecious, inflorescences terminal, paniculate, long-lived, with large, overlapping, tubular bracts [c.f. Alpinia sect. Myriocrater!], pedicels not articulated; C 3 + 3, connate; staminate flowers: nectariferous lobes opposite A, A 3, connate, extrorse, pollen ?tricolpate; carpellate flowers: G [8-13], ovules basal, micropyle bistomal, funicle well developed, nucellar cap +, stigmas sessile, ray-like, with an opening in the middle; fruit a fleshy capsule with lignified commissural valves; seeds with aril from funicle/outer integument; exotegmic fibers +; endosperm slight, perisperm +, very scanty; n = 9. 1/1: Scyphostegia borneensis. Borneo, not the southern part. [Photo - Flower, Leaf, Inflorescence.] - see Metcalfe 1954 (anatomy); van Heel 1967 (flowers and fruits); Hutchinson 1973 (the diversity of early interpretations of the gynoecium); Chase et al. 2002 (general)]).

LACISTEMATACEAE Martius, nom. cons.   Back to Malpighiales

Trees; plants Al accumulators; chemistry?; vessel elements with scalariform perforation plates; sieve tubes?; petiole bundle D or deeply C-shaped, also wing bundles +; leaves two-ranked, (entire); inflorescence a raceme to dense spike; flowers small, P cup-like [1-6], A 1, the thecae ± separated and even stipitate, G [2-3], median member adaxial, 1-2 ± apical ovules/carpel, funicles thick, long, style branches short, ?stigma; fruit a 1(-3)-seeded capsule; testa fleshy or not; endosperm copious, embryo (short), with foliaceous cotyledons; n = ?

2[list]/14. Greater Antilles (Jamaica), Mexico southwards, not in Chile. (Map: from Sleumer 1980).[Photo - Flower, Fruit]

• Lacistemataceae are woody, evergreen plants that have stipulate and usually serrate leaves. The small flowers are borne in rather dense racemes to spikes and have an inconspicuous perianth and a single anther whose two thecae are often well separated and even stipitate. The dehiscent fruit is small, with one, sometimes to three, apical seeds.

Lacistemataceae do not cluster with the rest of Salicaceae and Kiggelariaceae (Savolainen et al. 2000a; Chase et al. 2002), although they are probably in this area (Chase et al. 2002; see also D. Soltis et al. 1999, 2000). Davis et al. (2005a) place them as sister to Salicaceae s.l. (61% bootstrap, 100% posterior probability), as do Korotkova et al. (2007: slightly higher jacknife); as might be expected, they lack salicoid teeth.

Is there an aril in Lacistemataceae? Sleumer (1980) records one for Lacistema, but a fleshy seed coat for Lozania; an aril is obvious in neither. In the latter genus there appear to be long "hairs" inside the fruit which perhaps support the dangling seed; these hairs are thick-walled but unlignified cells that may be derived from the funicle.

See Sleumer (1980: as Flacourtiaceae - Lacistemeae) for a monograph. Lozania - Riviere 270 (anatomy), Gentry et al. 22231 (fruit); Lacistema - Aymard & Delgado 6882 (fruit), Rimachi Y. 11201 (anatomy - stomata tending to anisocytic).

Turneraceae + Malesherbiaceae + Passifloraceae [= Passifloraceae s.l.]: cyclopentenoid cyanogenic glycosides and/or cyclopentenyl fatty acids +, cyanogenic glycosides derived from valine and isoleucine + [?Malesherbiaceae]; (plant with unpleasant smell); (foliar glands +), colleters +; K + C together forming a tube, (corona or scales on C), A 5 [opposite sepals], styles +; aril +, endotestal cells large, exotegmen palisade, endotegmen persistent; endosperm persistent, oily.

The cyclopentenoid glycosides may be sequestered by caterpillars feeding on plants of these families and perhaps even used as nitrogen sources; Achariaceae also have this combination of features.

Turneraceae are weakly associated with Malesherbiaceae in Chase et al. (2002), the two being strongly associated with Passifloraceae. Turneraceae show biparental or paternal transmission of plastids, as may Passifloraceae (Shore et al. 1994). Including Turneraceae and Malesherbiaceae in Passifloraceae s.l. is an optional arrangement in A.P.G. II. However, basic clade limits within this group may need re-evaluating, since preliminary data suggest that a paraphyletic Passifloraceae may include Turneraceae and Malesherbiaceae (A.P.G. II 2003), while Korotkova et al. (2007: three taxa from the three families) found Turnera and Passiflora to be sister taxa with 98% jacknife support.

For a discussion on aril development in this group, see Kloos and Bouman (1980); although it is often described as funicular, they incline to call it raphal.

Turneraceae + Malesherbiaceae: leaves spiral; exotestal cells arranged in lines; x = 7.

TURNERACEAE Candolle, nom. cons.   Back to Malpighiales

Herbaceous or woody; plant with unpleasant smell; ellagic acid 0; cortical vascular bundles [= leaf traces] common; vessel elements with simple (and scalariform) perforation plates; stomata various; leaves conduplicate, glands often at base of lamina, stipules 0 (+ - e.g. Erblichia); bracteoles often large; (flowers heterostylous), C contorted, deliquescent, (glands or corona at mouth of tube), nectary near base of tube (on sepals; filaments), microsporogenesis simultaneous, G [2, 3] (half inferior), micropyle zig-zag, hypostase +, stigmas concave, often ± penicillate; K + C tube deciduous; aril +; seeds often regularly rugose; n also = 5 (13).

10[list]/205: Turnera (122), Piriquetia (44). Tropical to warm temperate America and Africa (inc. Madagascar and Rodriguez I.) (Map: from Wickens 1976; Heywood 2007). [Photo - Flower]

• Turneraceae are herbs to woody plants that may be recognised by their often very hairy, serrate, exstipulate leaves with strong secondary venation. The flowers, with their calyx and corolla together forming a tube, contorted and deliquescent yellow or sometimes red corolla, parietal placentation, and often bifid, fringed stigmas, are distinctive. The loculicidally capsular fruits have arillate seeds with a minutely and often regularly reticulate testa.

Turneraceae are the hosts of caterpillars of several genera of Nymphalidae, alternate hosts include Salicaceae, Passifloraceae, and Violaceae (Arbo 2006 and references). It has been suggested that there is floral mimicry between Turnera and Malvaceae in Argentina (Benitez-Vieyra et al. 2007). Species of both Turnera and Piriqueta have epiphyllous flowers. Heterostyly is common in these and some other genera of Turneraceae.

Some information is taken from Vijayaraghavan and Kaur (1967: embryology), Hegnauer (chemistry), González and Arbo (2005: anatomy), and Arbo (2006: general account).

Synonymy: Piriquetaceae Martynov

MALESHERBIACEAE D. Don, nom. cons.   Back to Malpighiales

Herbaceous or subwoody; hairs conspicuous, multiseriate, often glandular; plant with unpleasant smell; tannins?; (cork cortical); vessel elements usu. with simple perforation plates; nodes also 1:1; leaves often deeply lobed, (margins entire; stipules foliaceous or 0), colleters?; K + C tube long, K valvate, C valvate, (denticulate corona at the mouth of the tube), androgynophore +, G [3, 4], micropyle endostomal, large protrusion from chalazal region, styles slender, stigmas capitate-clavate, ?type; K + C tube persistent; seeds pitted, aril 0; endosperm type?

1[list]/24. Andean South America from Peru S., esp. N. Chile (Map: see Gengler-Novak 2002). [Photo - Habit]

• Malesherbiaceae are more or less herbaceous, foetid and often densely glandular-hairy plants with distinctive flowers. The calyx and corolla together form a long tube; the sepals and petals are both valvate. There is an androgynophore, the floral tube is persistent in fruit, and the styles are often subapical on the capsule.

For relationships within the family, see Gengler-Novak (2002, 2003).

Cronquist (1981) suggests that stipules are lacking. Malesherbiaceae are very close to Turneraceae, but their seeds lack arils, the corolla aestivation differs, etc.; they are also close to Passifloraceae (Chase et al. 2002). The styles are shown as being commissural by Schnizlein (1843-1870: fam. 198).

General information is taken from Ricardo S. (1967, he suggested that the micropyle is endostomal) and Kubitzki (2006b); for chemistry, see Hegnauer (1969, 1990).

PASSIFLORACEAE Kunth, nom. cons.   Back to Malpighiales

Woody or herbaceous climbers with simple branch tendrils; flavonols +, ellagic acid +/0, tannins 0; anomalous secondary thickening quite common; vessel elements with simple perforation plates; wood often fluorescing; supernumerary buds +; leaves spiral, (compound), conduplicate, (margins entire), 2ndary veins often palmate, glands common on petiole or lamina; (dioecious), inflorescence cymose; flowers (3-)5-merous, corona of (1-)2-several rows of filaments or membranes (0), nectary ± on K/C tube, (A basally connate), gynophore or androgynophore frequent, anthers versatile, pollen to 12-colporate, G [(2-)3(-7)], funicle often long, nucellar apex pointed, micropyle bistomal or zig-zag, stigmas capitate or divided [Adenia], papillae multicellular; fruit usu. a berry; seeds often flattened, sculpted, bony, hairy or not, ruminate, aril +, testa multiplicative, sarcoexotestal or not, endotesta crystalliferous, lignified or not; n = 6 (7) 9(-12).

16[list]/705 (10/675): Passiflora (525, not Africa, inc. Hollrungia and Tetrapathea - Krosnick & Freudenstein 2006), Adenia (100: see Hearn 2006 for a phylogeny, great variation in life form). Tropics to warm temperate, especially Africa and America (Map: from van Balgooy 1975; George 1982). [Photo - Collection]

Paropsieae

Trees or shrubs; vessel elements in multiples, with scalariform perforation plates; leaves spiral, reduced [orthotropic axes], or two-ranked [plagiotropic axes], 2ndry venation pinnate, (stipules 0); inflorescence racemose; (A -20), androgynophore +/0, (pollen 6-porate), nectary 0 (annular), G [(2-)3-6], (style + [Barteria]); seeds not sculpted; n = ?

6/ca 27: Paropsia 12. Tropical, esp. (west) Africa.

• Many Passifloraceae are tendrillar vines or lianes; the leaves are alternate, stipulate, often with palmate venation, and often with glands on the lamina or petiole. The tendril is often oblique to the axis; a flower may be lateral to it, or paired flowers are borne on it or at its base. The axillary buds are superposed, and the lower bud produces the inflorescence/tendril. Floral features are, however, more important for the recognition of the family. The calyx and corolla are often similar and fused to each other at the base, and there is usually a complex corona. There is a gynophore or androgynophore, often five stamens with large and versatile anthers, and a three-carpellate gynoecium with parietal placentation. The fruit is a berry, and the often flattened seeds are arillate.

Passiflora and its relatives are known for their association with Heliconius butterflies, and the former show great variation in leaf morphology, foliar glands (some of these are involved in egg mimicry - Vanderplank 2007 for references), etc. Heliconius itself is also closely associated with Anguria (Cucurbitaceae), which it pollinates, also obtaining nutrients from the pollen (e.g. Gilbert 1972, 1975; Spencer 1988). The larvae of some Acraeinae and also of brightly-coloured Notodontidae-Dioptininae moths are also often found on Passiflora (Miller 1992; Silva-Brandão et al. 2008), and at least the former are also found on Barteria. Details of the association between the African ant-plant Bartera fistulosa and the ant Tetraponera aethiops are given by Dejean et al. (2008).

Flacourtiaceae - Paropsieae (Barteria, Paropsia, etc.) are to be included here (Chase et al. 2002); I have characterised them separately above. Anatomically the two are rather similar, indeed, the major variation in the family seems to be associated with habit - lianes versus tress (Ayensu & Stern 1964). Passiflora and immediate relatives have stem collenchyma, cymose inflorescences, and branches developing from an accessory (superposed) bud, as is common in taxa with axillary tendrils with non-basal prophyllar buds, which in this case may be flowers. Adenia may be rather different from other Passifloraceae, perhaps more like Malesherbiaceae and Turneraceae, e.g. in moderately developed corona, tricolporate pollen (e.g. see Feuillet & MacDougal 2006). Indeed, Adenia has a nectary often made up of separate glands, a hollow style, and its stigma lacks multicellular papillae (Bernhard 1999a, c), in addition, it may be dioecious, it lacks an androgynophore, the stamens are sometimes connate, and there is a gynophore; some species have a true hypanthium (de Wilde 1971b)!

Cyanogenic glycosides in this family have a variety of precursors, both protein and non-protein amino acids (Miller et al. 2006 for refernces). In species of Passiflora with strongly bilobed leaves, ptyxis may be modified conduplicate: the blade makes a V with an inverted V at the end of each arm. The tendril is an axillary shoot and flowers can arise from prophyllar buds. Sazima and Sazima (1978) note that the bat-pollinated flowers of Passiflora mucronata become zygomorphic as the stamens move after the flowers opens; Endress and Matthews (2006a) give this as an example of monosymmetry in the family. For floral anatomy of Passiflora, see Puri (1947), for a general account of the genus, see Ulmer and MacDougal (2004), for a formal infrageneric classification, see Feuillet and Macdougal (2004), for floral development, see Krosnick et al. (2006), and for the phylogeny of Passiflora, see Yockteng and Nadot (2004), Krosnick and Freudenstein (2005: also morphology). Hansen et al. (2006) discuss chromosome number evolution, n = 12 may be the basal number; see also de Melo and Guerra (2003).

Do the sieve tubes have non-dispersive protein bodies?

Carpel orientation is taken from Le Maout and Decaisne (1868) and Schnizlein (1843-1870: fam. 197); for pollen, see Presting (1965) and Spirlet (1965), for chemistry, see Hegnauer (1969, 1990), for embryology, etc., see Singh (1970), for branching, see de Wilde (1971a), for general information, de Wilde (1974), for anatomy, see Harms (1893), for stipules, see Dahlgren and van Wyk (1988), and for a general account, see Feuillet and MacDougal (2006). I am grateful to J. M. MacDougal for information.

Synonymy: Modeccaceae Horaninow, Paropsiaceae Dumortier, Smeathmanniaceae Perleb

GOUPIACEAE Miers   Back to Malpighiales

Evergreen trees; plants Al-accumulators, otherwise chemistry unknown; vessel elements with scalariform perforations; nodes ?; petiole with ± annular bundle(s), plus inverted medullary bundles; branched sclereids +; cuticle waxes 0; leaves two-ranked, tooth ?type, 2ndary veins actinodromous, 3ary veins scalariform; inflorescences axillary, umbellate, pedicel articulation?; C induplicate-valvate, long, apical part inflexed, disc annular, connective shortly prolonged, with long hairs, pollen with endexinal folds, G opposite petals, placentation axile, several basal ovules/carpel, styles separate, short, on outer shoulders of carpels, stigmas subulate, type?; fruit a drupe; testa and tegmen ca 6 cells thick, testa with one layer [mesotestal] of sclereids, exotegmen poorly developed; endosperm copious; n = ?

1/2. N.E. South America.

• Goupiaceae may be recognised by their two-ranked stipulate leaves with ascending secondary veins and scalariform tertiaries. The inflorescence is umbellate and the petals are relatively long and inflexed at the apex. The gynoecium has basal-axile placentation and widely separated styles; the fruit is a single-seeded drupe.

The family is poorly known. Cronquist (1981) included Goupiaceae in Celastraceae, Takhtajan (1997) in Celastrales, A.-L. de Jussieu and others have placed it in Rhamnaceae. It is often suggested that only seedlings have dentate leaves, those of the adult being entire, but leaves of flowering specimens are in fact frequently toothed.

Information on pollen is taken from Lobreau-Callen (1980), and on seed, from Takhtajan (2000).

Lophopyxidaceae + Putranjivaceae: stomata paracytic; 2 apical epitropous ovules/carpel, style branches short or 0; fruit 1-seeded.

This clade may have separated in the Cretaceous-Albian 111-100 million years before present, Lophopyxidaceae and Putranjivaceae themselves diverging at end Coniacian or thereabouts ca 85 million years before present (Davis et al. 2005a).

Davis et al. (2005a) find a strong association between these two families; they may in turn be associated with the group of families with parietal placentation.

LOPHOPYXIDACEAE H. Pfeiffer   Back to Malpighiales

Liane with leaf tendrils; chemistry?; branches with lateral bud at base; secondary thickening with included phloem; vessel elements with simple perforation plates; phloem stratified; nodes ?; petiole bundles arcuate; leaves spiral; plant monoecious, inflorescence branched, flowers in clusters, sessile, small, K connate basally, valvate, C very small, staminate flowers: stamens = and opposite sepals, cordate glands adnate to C; carpellate flowers: glands forming a lobed disc, G [(4) 5], opposite petals, small funicular obturator +, stigmas subulate; fruit a 5-winged samara; seed coat?; endosperm ?development, +; n = ?

1/1: Lophopyxis maingayi. Malesia to the Solomon and Caroline Islands (Map: from Sleumer 1971b).

• Lophopyxidaceae are tendrillate lianas with quite strongly serrate leaves that have pinnate venation; the flowers are small and are borne in tight clusters along the branched inflorescences. The one-seeded fruit has five wings.

Included in Celastraceae by Cronquist (1981) and Hutchinson (1973), in Celastrales by Takhtajan (1997), but placed close to Pandaceae (represented by Microdesmis) by Savolainen et al. (2000a; see also Chase et al. 2002). Sleumer (1971b) described the tendrils as being leaves and also bud-bearing branches; the ultimate spirally-recurved portion seems to be foliar.

PUTRANJIVACEAE Endlicher   Back to Malpighiales

Evergreen trees; cucurbitacins [triterpenes], glucosinolates, biflavonoyls +; cork?; vessel elements with scalariform perforation plates; petiole bundles elliptic; hairs unicellular; leaves two-ranked, (veins running into opaque deciduous teeth, or spines); plant dioecious, inflorescence fasciculate; P 4-5(-7), staminate flowers: A (2-)3-20(-many; extrorse), disc + or 0; carpellate flowers: G 1[-4(-9)], ovules epitropous, nucellus ca 2 cells thick, disintegrating early, micropyle (exo/)endostomal, outer integument 3-8 cells and inner 6-14 cells across, archesporium 2-3-celled, endothelium +, placental obturator +, stigmas flap-like, ?type; fruit a drupe; testa vascularised, exomesotesta sclereidal, tegmen 6-24 or more cells thick, exotegmen cells cuboidal; endosperm copious; n = (19) 20 (21).

3[list]/210: Drypetes (200). Tropical, esp. Africa and Malesia (Map: from FloraBase 2005; Andrew Ford, pers. comm). [Photo - Flower, Fruit]

• Putranjivaceae are trees to shrubs that may be recognised by their two-ranked often rather coriaceous leaves that are asymmetrical at the base and dry a greyish color; they frequently taste peppery or like an inferior radish when fresh, although the taste may take a little time to develop (mustard oils!). The fasciculate flowers are frequently rather small and the single-seeded drupe is often crowned by the persistent, flap-like stigmas.

Perhaps not surprisingly, caterpillars of Pierid butterflies have quite often (23/2690 records) been recorded from this group (see also Brassicales and Fabaceae) - nothing so far known from Lophopyxidaceae!

Putranjivaceae have usually been included in Euphorbiaceae (as by Webster 1994, in Phyllanthoideae), but can be distinguished i.a. by their chemistry, embryology, and fruit. They are also certainly not to be placed with the rest of the glucosinolate families in Brassicales (e.g. Rodman et al. 1997, 1998).

For chemistry, see Hegnauer (1966, 1989, as Euphorbiaceae), for embryology and seed anatomy, see Singh (1970), Stuppy (1996), and Tokuoka and Tobe (1999, 2001 - Lingelsheimia included, but tegmen 3-4 cells thick and testa vascularised, to be placed in Phyllanthaceae - see Kathriarachchi et al. 2005), for wood anatomy, Hayden and Brandt (1984 - it is like that of Aporusa, etc. [= Phyllanthaceae]). For a checklist and bibliography, see Govaerts et al. (2000).

CTENOLOPHONACEAE Exell & Mendonça   Back to Malpighiales

Woody; ellagic acid?; vessel elements with scalariform perforation plates; oxalate crystals +; cuticle waxes 0; stomata anomo- or anisocytic; hairs tufted; petiole bundle arcuate; leaves entire; inflorescence terminal, ?thyrsoid; C contorted, caducous, disc +, A adnate to base of disc, pollen 3-9 zonocolporate, G [2], ovules with placental obturator, style +, branches short, stigmas capitate; fruit a ? capsule, K persistent, swollen; seed single, persisting on columella; arillode ± hairy, exotestal cells palisade, the outer wall thickened; endosperm copious, cotyledons very large, folded; n = ?.

1[list]/3. W. Africa, Malesia (Map: from van Hooren & Nooteboom 1988b).

• Ctenolophonaceae are trees that may be recognised by their opposite, entire leaves with interpetiolar stipules and their terminal inflorescences with rather elongated flower buds. The rounded sepals are persistent and accrescent, and the grey-drying and closely-ribbed fruit is distinctive. There is a single seed half enveloped by the hairy arillode; it remains attached to the slender central columella.

Ctenolophonaceae may have diverged in the Cretaceous-Albian 111-100 million years before present ([109.6-]101.8[-96.6]/[97.1-]91.0[-88.1] million years before present: Davis et al. 2005a). Fossil pollen is known from South America and India, and the first records are from Africa in the Upper Cretaceous (Muller 1981).

Like Humiriaceae, there are "marginal" stomata on the disc and the anthers have a broad connective (Link 1992b); the wood anatomy is also similar.

Some information is taken from van Hooren and Nooteboom (1988b).

Erythroxylaceae + Rhizophoraceae: tropane [hygroline] and pyrrolidine alkaloids, non-hydrolysable tannins +; sieve tube plastids with protein crystalloids; mucilage cells common; stomata paracytic; leaves involute, colleters +; inflorescence cymose; K valvate, C conduplicate, petals enclosing a stamen/stamens, filaments connate basally, median G adaxial, ovules epitropous, inner integument ca 6 cells thick, endothelium +; fruit a septicidal capsule, K persistent; seeds arillate, exotestal cells enlarged, thick-walled, ± tanniniferous; endosperm starchy, embryo green.

This clade may have diverged in the Cretaceous-Aptian around 114 million years before present ([119.3-]113.8[-110.2]/[105.7-]101.6[-102.1] million years before present: Davis et al. 2005a).

There is weak support for an association of [Caryocaraceae [Linaceae + Irvingiaceae}} with this clade (Soltis et al. 2007a), and there are a number of features in common, such as a basally connate androecium, epitropous ovules with an endothelium, etc. (Matthews & Endress 2007). Ctenolophonaceae, etc., might also be associated, but their floral similarities did not seem to be so great.

Including the two families in Rhizophoraceae s.l. is optional in A.P.G. II, but the two are so well known and for the most part distinctive that keeping them separate is preferable. Rhizophoraceae used to be placed in Myrtales (Cronquist 1981) or Myrtanae (Takhtajan 1997), largely because of their vestured pits and inferior ovary, but they are well supported as sister to Erythroxylaceae (e.g. Setoguchi et al. 1999; Schwarzbach & Ricklefs 2000; Chase et al. 2002; Korotkova et al. 2007). Although an unexpected family pair, when comparing Aneulophus (Erythroxylaceae) with non-mangrove Rhizophoraceae, the differences are then less obvious.

For floral development, see Matthews and Endress (2007).

ERYTHROXYLACEAE Kunth, nom. cons.   Back to Malpighiales

Smallish trees and shrubs; ellagic acid 0; vessel elements with simple perforation plates; wood commonly with SiO₂ grains; nodes with lateral bundles originating well before the central, forming cortical bundles; sclereids +; petiole bundle arcuate to annular with medullary and adaxial bundles; branching from previous flush; buds perulate; leaves also usu. two-ranked (spiral) and stipules intrapetiolar; inflorescence often fasciculate; (pedicel not articulated - Aneulophus?), heterostyly common, K connate basally, C with fringed bilobed ligule (0), obdiplostemonous, extrorse, (connective not thickened), pollen trinucleate, nectary 0, G [(2-)3(-4)], (adaxial only fertile), also 1 ovule/carpel, micropyle endostomal, style (short), (hollow), branches separate, stigmas ± capitate; fruit often a 1-seeded drupe, A also persistent; (endosperm 0); n = 12.

4[list]/240: Erythroxylum (230). Pantropical, esp. American (Map: from van Steenis and van Balgooy 1966; Heywood 1978). [Photo - Flower, Fruit.]

• Erythroxylaceae are shrubs to trees that can be recognised by their entire leaves, stipules that are at least in part interpetiolar, and usually fasciculate inflorescence with rather small flowers; both calyx and filaments persist at the base of the fruit. Erythroxylum itself has intrapetiolar stipules, there are often longitudinal markings on the leaves, the filaments are often of two different lengths and are basally connate, and the fruit is a drupe.

Cocaine is sequestered by the larvae of Eloria noyesii, a lymanitrid moth.

Aneulophus has a thick testa, thin tegmen, aril, opposite leaves, colleters, inter/intrapetiolar stipules, and a septicidal capsule; from petal length, it appears that the flowers are monosymmetric.

The leaves of Erythroxylum coca were described as being revolute by Cullen (1978); they are involute (e.g. Peyritsch 1878; Keller 1996). The nodes were described as being unilacunar by Sinnott (1914). Erythroxylum sometimes has milky exudate. Are the lamina teeth theoid??

For chemistry, see Hegnauer (1966, 1989) and Aniszewski (2007).

Synonymy: Nectaropetalaceae Exell and Mendonça

RHIZOPHORACEAE Persoon, nom. cons.   Back to Malpighiales

Trees; ellagic acid +; vessel elements with simple and/or scalariform perforation plates; true tracheids +; pits vestured; nodes also multilacunar, often with split-laterals; subepidermal laticifers in flower; cristarque cells?; branching from current flush; leaves (supervolute - mangroves), (margins entire); inflorescence axis often evident, pedicel articulation?; K (3-)4-5(-16), C small, often hairy, variously lobed, fringed, or with filiform appendages, aristate, A 9-many, (fasciculate; free; connective not well developed), disc on ovary or hypanthium (0), G [2-many], opposite sepals, when 2, lateral, micropyle zig-zag or endostomal, inner integument 5-20 and outer integument 2-5 cells across, several megasporocytes, style + (branched - Gynotroches), stigma ± punctate, capitate or lobed, ?type; (endotesta crystalliferous); (embryo short), cotyledons large; n = (13), 14, 16, 18, 21; germination epigeal, cotyledonary node unilacunar.

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