MALPIGHIALES Martius Main Tree, Synapomorphies

Habit Bisexual, monoecious or dioecious (rarely andromonoecious, gynomonoecious, polygamomonoecious, androdioecious, or polygamodioecious), evergreen or deciduous trees, shrubs, suffrutices or lianas, perennial, biennial or annual herbs (sometimes climbing).

Vegetative anatomy Phellogen ab initio epidermal or subepidermal, pericyclic, or outer- or inner-cortical. Secondary lateral growth normal, anomalous (from cylindrical cambium or several concentric cambia) or absent. Vessel elements with simple or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, scalariform or opposite, simple or bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate (often also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (rarely homocellular), or absent. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, vasicentric, aliform, lozenge-aliform, winged-aliform, confluent, scalariform, reticulate, unilateral, or banded, or absent. Intraxylary phloem sometimes present. Sieve tube plastids Ss, S₀, Pcs or Pc type. Nodes 1:1, unilacunar with one leaf trace, 3:3, trilacunar with three traces, or ≥5:≥5, multilacunar with five or more traces. Schizogenous secretory cells, canals or cavities or glands with resin, balsam or other secretions often abundant. Laticifers sometimes present. Heartwood often with gum-like substances. Silica bodies sometimes present. Cristarque cells sometimes present. Calciumoxalate as prismatic, rhomboidal or acicular crystals, crystal sand, druses, styloids or other types.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, furcate, T-shaped, malpighiaceous hairs, stellate, candelabra-shaped, dendritic, lepidote or peltate, or absent; glands and glandular hairs sometimes present; stinging hairs occasionally present.

Leaves Usually alternate (spiral or distichous) or opposite (rarely verticillate), pinnately or palmately compound, or simple and entire or lobed, with conduplicate, supervolute, convolute, involute, revolute, curved or flat ptyxis (rarely absent). Stipules interpetiolar, intrapetiolar or petiolar, free or connate, often rudimentary or absent (sometimes modified into spines or hair-like); leaf sheath absent. Petiole often articulated, sometimes geniculate. Petiole vascular bundle transection arcuate, annular or complex; petiole sometimes with lateral flank bundles. Venation pinnate or palmate, eucamptodromous, brochidodromous, reticulodromous or parallelodromous (rarely acrodromous, actinodromous or campylodromous). Stomata anomocytic, paracytic or anisocytic (rarely cyclocytic or tetracytic). Cuticular wax crystalloids usually as rosettes of platelets (Fabales type; sometimes absent). Domatia as pits or hair tufts (rarely myrmecodomatia). Epidermis with or without mucilaginous idioblasts. Mesophyll often with sclerenchymatous idioblasts. Leaf margin or leaflet margins entire, crenate or serrate; teeth with one vein proceeding into congested caducous tooth apex; glandular teeth, salicoid or violoid, rarely present on leaf margin.

Inflorescence Terminal or axillary, panicle, fascicle, thyrsoid, corymb, raceme-, spike-, catkin- or umbel-like cymose, or racemes, spikes or catkins (sometimes pseudanthium), or flowers solitary axillary. Bracts and/or floral prophylls (bracteoles) sometimes absent.

Flowers Actinomorphic or zygomorphic. Pedicel often articulated. Hypanthium rarely present. Usually hypogyny (rarely epigyny or half epigyny). Sepals (two or) three to five (to c. 20), with imbricate, valvate, decussate or open (sometimes truncate) aestivation, usually whorled (rarely spiral and indistinctly separate from petals), free or connate at base. Petals (two to) four to 15, usually whorled (rarely spiral and indistinctly separate from sepals), with imbricate, valvate, contorted, involute, decussate or crumpled (rarely cochlear or open) aestivation, usually clawed, usually free (sometimes more or less connate, or absent). Corona sometimes present at petal bases. Nectaries at filament bases or absent. Disc slightly developed or absent (nectariferous disc sometimes extrastaminal, lobate or cupular, usually annular).

Androecium Stamens one to more than 750, usually in one or more whorls. Filaments free or more or less connate, often in one or three fascicles, usually free from tepals (sometimes epipetalous). Anthers basifixed or dorsifixed, usually non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse (sometimes extrorse, rarely latrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum usually secretory (rarely amoeboid-periplasmodial). Staminodia one to more than 50, extrastaminal or intrastaminal, or absent.

Pollen grains Microsporogenesis usually simultaneous (rarely successive). Pollen grains (2–)3(–6)-colpor(oid)ate, tri- to pentacolp(oid)ate or tetra- to polyporate (rarely syncolpate or inaperturate), usually shed as monads (sometimes tetrads), usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate (rarely intectate), with columellate or granular infratectum, perforate, reticulate, microreticulate, or striate, rugulate, fossulate, foveolate, scabrate, verrucate, spinulate, echinate, retipilate, psilate or smooth.

Gynoecium Pistil composed of two to ten (to 20) usually connate carpels. Ovary usually superior (rarely inferior or semi-inferior), unilocular to quinquelocular (to 20-locular). Style single, simple, or stylodia two to five (to more than twelve), usually free, or absent; style sometimes unifacial. Stigma one, capitate to peltate, or stigmas two to five, capitate, punctate or truncate, papillate, usually Dry (sometimes Wet) type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation axile, apical, subbasal or parietal (rarely laminar). Ovules one to more than 50 per carpel, anatropous or hemianatropous (sometimes amphitropous, orthotropous, or anacampylotropous), ascending, horizontal or pendulous, apotropous or epitropous, usually bitegmic, usually weakly crassinucellar or incompletely tenuinucellar. Micropyle bistomal or endostomal (sometimes exostomal). Funicular or placental obturator sometimes present. Archespore usually unicellular (rarely bicellular or tricellular). Nucellar cap or nucellar beak sometimes present. Megagametophyte usually monosporous, Polygonum type (sometimes tetrasporous, 16-nucleate, Penaea type, or disporous, Allium type, etc.). Antipodal cells sometimes proliferating, sometimes absent. Endosperm development ab initio nuclear. Endosperm haustoria chalazal or absent. Embryogenesis usually solanad (sometimes onagrad or asterad, rarely piperad or caryophyllad).

Fruit A loculicidal and/or septicidal capsule, berry, drupe, nut, or schizocarp (divided into two to five nut-like, samaroid or drupaceous mericarps; rarely a secondary syncarp).

Seeds Aril or carunculus sometimes present. Seed coat testal, exotegmic or endotegmic (usually exotegmic). Testa sometimes vascularized, sometimes multiplicative. Sarcotesta sometimes present. Exotesta sometimes palisade. Mesotesta and/or endotesta sometimes lignified and/or sclerenchymatous. Tegmen sometimes multiplicative. Exotegmen and/or endotegmen often fibrous or lignified (sometimes palisade). Perisperm not developed. Endosperm copious or sparse, oily, or absent. Embryo large or small, straight or curved (rarely hook-shaped, spirally twisted or circinate), usually well differentiated, oily, with or without chlorophyll. Cotyledons two (to four). Germination phanerocotylar or cryptocotylar.

DNA Plastid gene rps16 often entirely or partially(lost in Passifloraceae, Violaceae, Salicaceae and Turneraceae, and also in some Linaceae and Malpighiaceae. Plastid gene atpF lost several times. Plastid gene infA lost/defunct. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, flavone-C-glycosides, afzelechin, biflavonoids, biflavanoids, biflavonoyls, trihydroxyflavonoids, flavonoid sulphates, cyanidin, cucurbitacins and other triterpenes, dammaranes, phorbole ester diterpenes, oleanolic acid derivatives, ellagic acid, methylated ellagic acids, gallic acid, non-hydrolyzable tannins (ellagitannins: geraniin, mallotussinic acid), condensed tannins, tannins with proanthocyanidins and catechin, proanthocyanidins (prodelphinidins), p-coumaric acid, caffeic acid, chlorogenic acid, cinnamic acid derivatives, tropane (hygrolinic) alkaloids (tropane-3α and tropane-3β-ols, tropacocaine, scopolamine oxides, hydroxytropines, teloidines, ecgonines, norecgonines, phyllalbine, oxytrapanes, brugine, etc.), indole alkaloids, pyrrolizidine alkaloids and other alkaloids (securinine, phyllantine, phyllochrisine, etc.), triterpene saponins, tyrosine-derived cyanogenic compounds, phenol glycosides (salicin, populin etc.), cyclopentenoid (cyclopentenylic) cyanogenic glycosides (gynocardin) and/or cyclopentenylic fatty acids, cyclopentenoid cyanhydrin glycosides derived from non-protein amino acid 2-(2-cyclopentenyl)glycine (in families near Achariaceae), xanthones (euxanthone, bixanthones, macluraxanthone, mangiferin, norathyriol, and anthraquinone xanthones), polyacetate-derived anthraquinones and arthroquinones, anthraquinones (vismiones etc.), hypericin, pseudohypericin, arbutin, emodin derivatives, biemodyles and closely allied compounds, benzophenones, acetophenones, anthrones, naphthodianthrones, coumarin derivatives substituted at position 4, syringaresinol, ferulic acid, phytosterols (sitosterol, stigmasterol), ethereal oils, hyperforin, picrotoxans, myo-inisitol, and nigracin present. Glucosinolates, benzylisoquinoline alkaloids and fluoroacetic acid rare.

The phylogeny within Malpighiales is highly unresolved (Korotkova & al. 2009; Wurdack & Davis 2009). Soltis & al. (2011) present a fully resolved tree, yet with most of the basal nodes weakly supported.

The clade [Achariaceae+[[Violaceae+[Passifloraceae+[Malesherbiaceae+Turneraceae]]]+ [Lacistemataceae+Salicaceae]+Goupiaceae]] have the following potential synapomorphies (Stevens 2001 onwards): ray cells crystalliferous; sieve tubes with non-dispersive protein bodies; cuticular waxes usually absent; pedicels articulated; nectariferous tissue present; stamens as many as sepals, antesepalous; median carpel abaxial; placentation parietal, with raised placentae; aril present; endotegmen persistent; and endosperm oily, persistent. Achariaceae, Malesherbiaceae, Turneraceae, and Passifloraceae often have some kind of corona or scales on the petals; and cyclopentenoid cyanogenic glycosides and/or cyclopentenylic fatty acids.

The clade [Passifloraceae+[Turneraceae+Malesherbiaceae]] has the following advanced features (Stevens 2001 onwards): leaf teeth with vein proceeding to opaque caducous apex; presence of colleters; sepals and petals together forming tube; stamens five, antesepalous; stylodia well developed; presence of funicular aril; endotestal cells large; exotegmen palisade; endotegmen persistent; and presence of cyclopentenoid cyanogenic glycosides and/or cyclopentenyl fatty acids. Turneraceae and Malesherbiaceae share the characters: leaves spiral; micropyle bistomal; exotestal cells arranged in lines; and x = 7. Moreover, Lacistemataceae and Salicaceae share the synapomorphies: small flowers; anthers ellipsoid to subglobose; and copious endosperm.

Lophopyxidaceae and Putranjivaceae have the following characteristics in common: stomata paracytic; stylar branches short or absent; placentation apical; ovules two per carpel; and fruit single-seeded.

The clade [Ctenolophonaceae+[Erythroxylaceae+Rhizophoraceae]] is characterized by: opposite leaves; stipules interpetiolar, enclosing terminal bud; articulated pedicels; extrastaminal nectary; ten stamens, of two different lengths; basifixed anthers, connate at base; postgenitally fused carpels; capitate to lobate stigmas, papillate; placentation apical; ovules two per carpel, collateral, pendulous, epitropous; megasporangium laterally thin, disintegrating; presence of endothelium; presence of placental obturator; sepals persistent in fruit; seed coat also exotestal; and presence of endosperm.

Erythroxylaceae and Rhizophoraceae share the synapomorphies, according to Stevens (2001 onwards): sieve tube plastids with protein crystalloids; presence of abundant mucilage cells; leaves with involute ptyxis; presence of colleters; stomata paracytic; inflorescence cymose; sepals with valvate aestivation, postgenitally fused; petals clawed, with conduplicate aestivation, enclosing stamen(s); antepetalous stamens longer than antesepalous stamens; median carpel adaxial; style somewhat impressed; inner integument approx. six cell layers thick; fruit a septicidal capsule; presence of aril; exotestal cells enlarged, thick-walled, tanniniferous; endosperm starchy; embryo with chlorophyll; presence of tropane (hygroline) and pyrrolidine alkaloids; and presence of non-hydrolyzable tannins.

The clade [[Clusiaceae+Bonnetiaceae]+[Calophyllaceae+[Hypericaceae+Podostemaceae]]] is characterized by: vessel elements with simple perforation plates; nodes unilacunar with one leaf trace; presence of secretory ducts; presence of schizogenous cavities; leaves with colleters; absence of stipules; stomata paracytic; leaf margin entire; inflorescence cymose; petals with contorted aestivation; absence of nectary; stamens numerous, often fasciculate, with antepetalous fascicles; carpels antesepalous or median carpel adaxial; stigma papillate; ovules numerous per carpel; micropyle bistomal; fruit a septicidal or septifragal capsule; anticlinal exotegmic cell walls sinuous, low, lignified; endosperm scarce or absent; embryo fusiform; and presence of flavonols, flavones, biflavonoids, and abundant xanthones. The clade [Calophyllaceae+[Hypericaceae+Podoste-maceae]] has leaves with gland dots or lines.

The clade [Centroplacaceae+[Elatinaceae+Malpighiaceae]] has fruits with persistent sepals. Malpighiaceae and Elatinaceae have the following potential synapomorphies in common: vessel elements with simple perforation plates; sieve tube plastids without starch and protein inclusions; leaves opposite (or verticillate); inflorescence cymose; flowers inverted; absence of nectary; when three carpels, then median carpel adaxial; fruit a septifragal capsule, with persistent calyx; endosperm scarce; and x = 6.

Ochnaceae, Medusagynaceae and Quiinaceae share the characters: presence of vestured pits; presence of cristarque cells; presence of mucilage cells and/or mucilage canals; leaves with secondary and tertiary venation well developed; petals with contorted aestivation; absence of nectary; and ovules tenuinucellate. Medusagyne and Quiinaceae have: separate styloids; well developed ovary roof; expanded stigma; and often two ovules per carpel. Moreover, Ochnaceae and Medusagyne have medullary vascular bundles.

The clade [Peraceae+[Rafflesiaceae+Euphorbiaceae]] has the following potential synapomorphies (Stevens 2001 onwards): vessel elements with simple perforation plates; flowers small, unisexual; carpels three; placentation apical; ovule one per carpel, pendulous, epitropous; presence of nucellar cap (unknown in Peraceae); stylodia separate; fruit a septicidal capsule or schizocarp, also splitting from columella and loculicidally; exocarp/mesocarp often separating from endocarp; seeds large; presence of micropylar carunculus; cotyledons longer and wider than radicula.

Phyllanthaceae and Picrodendraceae share the following synapomorphies: plant monoecious; stomata paracytic; flowers small; presence of style; placentation apical; ovules two per carpel, apical, epitropous; micropyle bistomal; parietal tissue at least ten cell layers thick; presence of obturator and nucellar beak; fruit an explosively dehiscent capsule, with fruit walls also splitting from persistent columella; exocarp/mesocarp often separating from endocarp; and x = 13.

The clade [Balanopaceae+[[Trigoniaceae+Dichapetalaceae]+[Chrysobalanaceae+Euphro-niaceae]]] is characterized by: hairs simple; ovules two per carpel, collateral; micropyle bistomal; outer and inner integuments at least five cell layers thick each; megasporangium evanescent by maturity; presence of endothelium; and endosperm scarce or absence. The clade [[Trigoniaceae+Dichapetalaceae]+[Chrysobalanaceae+Euphroniaceae]] has the following potential synapomorphies, according to Stevens (2001 onwards): vessel elements with simple perforation plates; presence of vestured pits; presence of mucilage cells; stomata paracytic; leaf margin entire; flowers obliquely zygomorphic; pedicels articulated; presence of hypanthium; sepals congenitally connate at base, with quincuncial aestivation, of unequal size (two outer sepals shorter), with epidermal mucilage cells; fertile stamens abaxial, connate; anthers much shorter than filaments, extremely introrse, with thecae almost in one plane; connective well developed abaxially with endothecium continuous over dorsal side of connective; presence of dorsal anther pit where filament joins; staminodia adaxial, absent in posterior most antepetalous position; gynoecium completely syncarpous up to stigma; carpel flanks slightly bulged out transversely, carpels thus demarcated from each other by longitudinal furrow; gynoecium and other floral parts with dense unlignified unicellular hairs; presence of style; stigma commissural; ovules epitropous, tenuinucellar; micropyle Z-shaped (zig-zag); outer integument two to five cell layers thick; inner integument three to eight cell layers thick; and presence of obturator.

Trigoniaceae and Dichapetalaceae share the characters: petiole vascular bundle transection arcuate; secondary veins strongly looping; inflorescence cymose; presence of mucilage cells in mesophyll of sepals (in addition to epidermis); nectary semi-annular, with lobes or scales; ovary and lower parts of style synascidiate; outer integument at most five cell layers thick; and testa multiplicative. Chrysobalanaceae and Euphroniaceae have the following features in common: presence of spurred hypanthium; petals clawed, with lignified hairs; and nectary present on adaxial side of hypanthium, usually annular, without lobes or scales.

Harms in Engler et Prantl, Nat. Pflanzenfam., Nachtr. 1: 256. Oct 1897, nom. cons.

Pangiaceae (Endl.) Blume in J. K. Hasskarl, Cat. Hort. Bot. Bogor.: 186. Oct 1844; Erythrospermaceae (DC.) Doweld, Tent. Syst. Plant. Vasc.: xxxii. 23 Dec 2001

Habit Usually dioecious (rarely monoecious), evergreen trees or shrubs (Acharieae consist of climbing herbs).

Vegetative anatomy Phellogen (in Lindackeria) ab initio superficial. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, scalariform or opposite, simple or reduced bordered pits. Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma absent (or very rare). Tyloses sometimes abundant. Sieve tubes with non-dispersive protein bodies?; sieve tube plastids S type. Nodes? Resinous substances etc. sometimes present in heartwood. Silica bodies present in some species. Acicular crystals and/or crystal sand present in some species. Prismatic crystals frequent; druses sometimes present.

Trichomes Hairs unicellular or multicellular, simple, stellate, peltate etc.

Leaves Alternate (spiral or distichous), simple, entire or lobate, with ? ptyxis. Stipules caducous (sometimes absent); leaf sheath absent. Petiole pulvinate, often geniculate. Petiole vascular bundle transection annular, with two lateral/adaxial bundles (in Lindackeria as inverted medullary plate). Venation pinnate. Stomata anomocytic, paracytic or anisocytic. Cuticular wax crystalloids? Domatia present in some species. Mesophyll sometimes with sclerenchymatous idioblasts. Leaf margin usually entire (sometimes serrate or crenate; salicoid teeth absent).

Inflorescence Terminal? or axillary, fasciculate, spike-like cymose, or racemose to spicate, or flowers solitary axillary.

Flowers Actinomorphic. Hypogyny. Sepals two to five, with open to valvate (Acharieae) aestivation, in one or two whorls or spiral, free or connate at base. Petals four to 15, with valvate (Acharieae) aestivation, in one or two whorls or spiral, usually free (in Acharieae three or four, connate into tube; rarely absent). Some genera with corona of scales, hairs or lobes at petal bases. Nectary usually absent. Disc absent.

Androecium Stamens three to numerous, antesepalous, antepetalous or irregular, in one or more whorls or groups, centripetally or almost synchronously developing. Filaments free; free from or more or less adnate to petals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; in Chiangiodendron and Kiggelaria poricidal, locellate, dehiscing by apical pores); connective sometimes widened. Tapetum secretory? Staminodia three to five, intrastaminal, or absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolpor(oid)ate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or microreticulate, often verrucate or psilate.

Gynoecium Pistil composed of two to ten usually connate carpels (in Erythrospermeae secondarily free). Ovary superior, unilocular. Style single, usually long (rarely short), sometimes branched. Stigma one or two to five, capitate to peltate, type? Pistillodium absent.

Ovules Placentation parietal. Ovules usually numerous (sometimes three to c. 20) per ovary (rarely one per carpel), usually anatropous (rarely orthotropous), bitegmic, crassinucellar. Micropyle endostomal or bistomal (in Acharieae, Z-shaped, zig-zag). Outer integument five or six cell layers thick, sometimes lobate. Inner integument five or six cell layers thick. Archespore usually unicellular (in Caloncoba bicellular). Nucellar cap and epistase present. Megagametophyte usually monosporous, Polygonum type (in Acharieae disporous, 8-nucleate), penetrating chalaza and forming caecum below tracheid ring. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril present or absent. Seed coat thick, usually pachychalazal. Testa usually distinctly vascularized (Acharieae lack vascular bundles in testa), in Acharieae with stomata. Exotestal cells elongate, sclereidal. Sarcotesta present (in i.a. Acharieae) or absent. Inner mesotesta sometimes sclereidal. Endotesta lignified, with sclereidal cells (sometimes radially elongate). Exotegmen usually non-fibrous, lignified (in Acharieae and Erythrospermum fibrous). Endotegmen? Perisperm not developed. Endosperm copious, oily. Suspensor absent. Embryo small, straight, well differentiated, with chlorophyll. Cotyledons two. Germination?

Phytochemistry Insufficiently known. Ellagic acid (in Kiggelaria) and cyclopentenoid (cyclopentenyl) cyanogenic glycosides (gynocardin) and/or cyclopentenyl fatty acids present.

Bentham et Hooker, Gen. Plant. 3: v, 341. 7 Feb 1880 [’Balanopseae’], nom. cons.

Distribution Queensland, Melanesia, with their largest diversity in New Caledonia.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with usually scalariform to reticulate (rarely simple) perforation plates; lateral pits alternate to almost opposite, simple pits. Vestured pits absent. Imperforate tracheary xylem elements fibre tracheids or thick-walled inconclusive libriform fibres with bordered pits. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually apotracheal diffuse, or paratracheal scanty. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Bark (and medulla?) with sclereids and rhomboidal crystals. Cortex with cristarque cells present in some species. Silica bodies present.

Trichomes Hairs unicellular, simple, often caducous; glandular hairs absent.

Leaves Alternate (spiral), simple, entire, coriaceous, with ? ptyxis. Stipules minute; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata usually anomocytic or laterocytic (sometimes cyclocytic). Cuticular waxes usually absent (crystalloids sometimes present as platelets). Calciumoxalate crystals? Leaf margin serrate; leaf teeth several cell layers thick, cells filled with dark tannin-like content.

Inflorescence Male flowers in axillary catkin-like cymose inflorescence; male flower with one bract. Female flowers solitary, surrounded by cupule-like organ consisting of spiral bracts with imbricate aestivation.

Flowers Actinomorphic, small. Hypogyny. Tepals in male flowers as small rudimentary teeth; female flowers without tepals. Nectary absent. Disc absent.

Androecium Stamens (one to) three to six (to 14). Filaments very short, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, latrorse to introrse (latero-introrse), longicidal (dehiscing by longitudinal slits); connective sometimes slightly prolonged. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tri- to pentacolp(oid)ate, shed as monads, bicellular at dispersal. Exine tectate, with columellate-granular infratectum, microperforate, beset with small spinules.

Gynoecium Pistil composed of (two or) three connate carpels. Ovary superior, (bilocular or) trilocular, often incompletely septate. Stylodia (two or) three, free or connate at base, once or twice bifid. Stigmas adaxial, long, type? Male flowers often with pistillodium.

Ovules Placentation subbasal. Ovules two per carpel, anatropous, intermediate between epitropous and apotropous, ascending, at least partially apotropous, bitegmic, weakly crassinucellar. Micropyle bistomal. Outer integument five to seven cell layers thick. Inner integument five to nine cell layers thick. Obturator absent. Parietal tissue approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A drupe with two or three single-seeded pyrenes and surrounded in lower part by cupule consisting of bracts.

Seeds Aril absent. Testa vascularized, persistent, with slightly thickened cell walls. Perisperm not developed. Endosperm sparse, thin. Embryo large, straight, with chlorophyll. Cotyledons two, cordate. Hypocotyl elongate. Germination phanerocotylar.

Phytochemistry Insufficiently known. Tannins and triterpenes abundant (especially in bark). Alkaloids not known. Ellagic acid?

Vegetative anatomy Phellogen ab initio superfical (in roots cortical). Vessels with simple or simple/transverse perforation plates; lateral pits alternate. Imperforate tracheary xylem elements usually thick-walled tracheids with simple pits?, non-septate? Wood rays uniseriate or multiseriate, heterocellular? Axial parenchyma paratracheal scanty (or apotracheal?). Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace (Archytaea, Ploiarium), or 3:3, trilacunar with three traces (Bonnetia). Mucilage cells frequent. Crystals?

Leaves Alternate (spiral), simple, entire, with involute or supervolute ptyxis. Stipules and leaf sheath absent. Colleters present in leaf axils. Petiole vascular bundle transection arcuate (Ploiarium) or complex. Venation pinnate, eucamptodromous, brochidodromous or parallelodromous; secondary veins ascending. Stomata paracytic. Cuticular wax crystalloids as rosettes. Epidermis in Bonnetia with mucilaginous idioblasts. Leaves and bracts in Archytaea and Ploiarium with vascularized disciform structures, absent in Bonnetia. Endodermis present. Mesophyll in Bonnetia with sclerenchymatous idioblasts; sclereids present (Bonnetia) or absent (Archytaea, Ploiarium). Leaf margin usually finely serrate (in young leaves with setae, associated with vascular tissue in Archytaea and Ploiarium, not in Bonnetia).

Inflorescence Axillary, cymose panicle or raceme- to umbel-like, or flowers solitary axillary (receptacle in Bonnetia ahogadoi developing into stolon with adventitious roots).

Flowers Actinomorphic. Hypogyny. Sepals five, with imbricate quincuncial aestivation, unequal, caducous or persistent, free, in Bonnetia and Ploiarium with apical bristle. Petals five, with contorted aestivation, free. Buds with long-pointed apex. Nectary and disc probably absent.

Androecium Stamens c. 40 to more than 100. Filaments thin, free or connate in one group at base (Bonnetia) or in five alternisepalous, antepetalous fascicles (Archytaea, Ploiarium), free from petals (Bonnetia) or adnate at base to petals (Archytaea, Ploiarium). Anthers basifixed to somewhat dorsifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Fasciclodium present or absent. Tapetum secretory? Staminodia five in Archytaea and Ploiarium, absent in Bonnetia.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate (rarely syncolpate), shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate? infratectum, finely reticulate.

Gynoecium Pistil composed of three to five connate carpels. Ovary superior, tri- or quadrilocular (Bonnetia), or quadri- or quinquelocular (Archytaea, Ploiarium). Style single, simple or trilobate to quinquelobate, or stylodia three to five, free. Stigmas papillate, type? Pistillodium absent.

Ovules Placentation axile. Ovules c. 30 to c. 50 per carpel, anatropous, pendulous or horizontal, bitegmic, tenuinucellar. Micropyle exostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Hypostase absent. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule (in Archytaea, Ploiarium dehiscing from proximal end; in Bonnetia dehiscing from distal end), usually with persistent central columella.

Seeds Aril absent. Exotestal cells thin-walled. Endotestal cells lignified (in Ploiarium elongate). Exotegmen with sinuous anticlinal cell walls? Endotegmen? Perisperm not developed. Endosperm sparse or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, small. Germination?

Phytochemistry Very insufficiently known. Xanthones (euxanthone, in Ploiarium bixanthones and anthraquinone xanthones) present.

Mesuaceae Bercht. et J. Presl, Přir. Rostlin: 218. Jan-Apr 1820 [’Mesuae’]; Cambogiaceae Horan., Prim. Lin. Syst. Nat.: 98. 2 Nov 1834 [’Cambogiaceae (Guttiferae)’]

Fossils Fossil pollen grains, Kielmeyeropollenites, are known from the Eocene of India. Symphonioxylon, fossil wood from Cretaceous and Miocene layers, may be ascribed to Calophyllaceae or Clusiaceae.

Habit Usually bisexual (occasionally cryptic-dioecious, rarely andromonoecious), usually evergreen trees (sometimes shrubs or epiphytes).

Vegetative anatomy Phellogen superficial or deeply seated. Endodermis in Kielmeyera often significant. Secondary lateral growth usually normal (in Endodesmia anomalous?). Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, usually simple pits. Vestured pits often present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with usually simple (rarely bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal or paratracheal. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Schizogenous secretory ducts or cavities or glands with resin, balsam or yellow to red secretions abundant (also in cortex and medulla). Colleters absent. Wood ray cells sometimes with silica. Crystals?

Trichomes Hairs unicellular or multicellular or absent (Caraipa and Marila with stellate hairs; Marila with branched hairs).

Leaves Alternate (spiral or distichous) or opposite, simple, entire, often coriaceous, usually with flat or conduplicate (in Kielmeyera supervolute) ptyxis. Stipules and leaf sheath absent. Paired modified colleters, “stipular glands”, sometimes present. Petiole bundle transection arcuate, annular or complex. Venation pinnate, eucamptodromous, brochidodromous or reticulodromous; tertiary venation sometimes scalariform or absent. Stomata paracytic. Cuticular wax crystalloids as rosettes. Lamina usually gland-dotted and/or with schizogenous secretory cavities (sometimes canals) with resin, balsam or yellow to red secretions (pellucid-punctate dots, resin/latex cavities). Leaf margin entire.

Inflorescence Terminal or axillary, cymose, often thyrsoid (sometimes racemose), or flowers sometimes solitary. Floral prophylls (bracteoles) sometimes (i.a. in Calophyllum and Lebrunia) absent.

Flowers Usually actinomorphic (in Marila asymmetralis obliquely zygomorphic), often large. Hypogyny. Sepals (two to) four or five (to c. 20), with imbricate quincuncial or decussate aestivation, usually free (rarely connate at base). Petals (three or) four or five (to eight), with contorted or decussate aestivation, free (absent in Calophyllum). Nectariferous disc usually absent (sometimes as separate units).

Androecium Stamens (four to) c. 20 to more than 100, not in distinct fascicles. Filaments usually in five alternisepalous, antepetalous indistinct groups, usually free (rarely connate), free from tepals. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, usually introrse (sometimes extrorse), usually longicidal (dehiscing by longitudinal slits, rarely poricidal, dehiscing by pores); connective often with small single or paired, sometimes apical, complex or simple glands (sometimes large and crateriform). Tapetum secretory. Staminodia two to more than 50, extrastaminal, or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely triporate or with several apertures), usually shed as monads (in Kielmeyera often as tetrads), bicellular at dispersal. Exine tectate or semitectate, with columellate? infratectum, reticulate, rugulate, fossulate, foveolate, scabrate or psilate.

Gynoecium Pistil composed of two to five connate antesepalous carpels (rarely monocarpellate?). Ovary superior, bilocular to quinquelocular. Stylodia two to five, free, usually long (longer than ovary), or style single, simple. Stigmas one to five, expanded to punctate, usually non-papillate, Wet type. Pistillodium?

Ovules Placentation usually axile (in Endodesmia clade apical, in Calophyllum and Kayea basal, in Clusiella laminar). Ovules usually two to numerous (in, e.g., Calophyllum, Endodesmia and Lebrunia one) per carpel, usually anatropous, ascending or horizontal, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad. Polyembryony present at least in Calophyllum and Kayea.

Fruit Usually a septicidal or septifragal capsule (in Kayea often with persistent and strongly accrescent calyx; sometimes a drupe; in Calophyllum, Clusiella and Mammea a berry).

Seeds Aril absent. Seed coat testal or exotegmic, sometimes winged. Testa with epidermis and exotegmen sinuous and with lignified cell walls, or testa multi-layered, complex and vascularized and exotegmen often absent. Endotegmen? Perisperm not developed. Endosperm sparse. Embryo small to large, straight (fusiform) or curved, rudimentary or well differentiated, with or without chlorophyll. Cotyledons two, massive, medium-sized to very large (Calophyllum, Mesua etc.; in Mammea connate). Germination phanerocotylar or cryptocotylar. Radicula in large-seeded speces often ephemeral and replaced by adventitious roots.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, biflavonoids, flavonoid sulphates, dammaranes, cyanidin, oleanolic acid derivatives, ellagic and gallic acids, proanthocyanidins (prodelphinidins), alkaloids, cyanogenic compounds isoprenylated xanthones (euxanthone, macluraxanthone, norathyriol), polyacetate-derived anthraquinones, coumarin derivatives substituted at position 4, and syringaresinol present.

Use Ornamental plants, fruits (Mammea americana etc), perfumes (Mammea siamensis), medicinal plants, cosmetics, dyeing substances, seed oils, timber.

Rhizobolaceae DC., Prodr. 1: 599. med Jan 1824 [’Rhizoboleae’], nom. illeg.; Rhizobolales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 60. Sep-Oct 1835 [‘Rhizoboleae’], nom. illeg.

Distribution Costa Rica to Paraguay and the West Indies, with their largest diversity in Amazonia.

Fossils Uncertain. Fossil pollen grains attributed to Caryocaraceae have been described from the mid-Eocene.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric. Tyloses abundant. Sieve tube plastids S type. Nodes ≥5:≥5, multilacunar with five or more leaf traces. Parenchyma with idioblasts containing branched sclereids (in medullary parenchyma) and solitary or groups of calciumoxalate crystals. Prismatic crystals often present.

Leaves Opposite (Caryocar) or alternate (Anthodiscus), bipinnate or trifoliolate to quinquefoliolate with articulated petiolules, coriaceous, with ? ptyxis. Stipules intrapetiolar (Anthodiscus) to interpetiolar, early caducous, or absent; leaf sheath absent. Stipulules often present (Caryocar), persistent or caducous. Colleters present. Petiole vascular bundle transection? Venation palmate (leaflet venation pinnate). Stomata usually anomocytic (sometimes anisocytic or paracytic). Cuticular wax crystalloids as smooth to irregular rosettes of platelets. Domatia as hair tufts. Epidermis with or without mucilaginous idioblasts. Mesophyll with sclerenchymatous idioblasts containing branched sclereids. Hydathodes usually present. Leaflet margins usually serrate (rarely almost entire).

Inflorescence Terminal, racemose or corymbose. Bracts usually absent (rarely present, small and caducous).

Flowers Actinomorphic, large (Caryocar) or medium-sized (Anthodiscus). Pedicel articulated at apex. Hypanthium sometimes present. Hypogyny. Sepals five (or six), with quincuncial to truncate or open aestivation, in Anthodiscus lobate, usually more or less connate. Petals five (or six), with quincuncial aestivation, caducous, free or connate in lower parts (Caryocar), or entirely connate forming caducous calyptra (Anthodiscus). Nectaries at base of filaments and ovary, or absent. Disc absent. Extrafloral nectaries often present on calyx.

Androecium Stamens 57 to at least 750, subperigynous. Filaments very long, often glandular-tuberculate at apex, connate at base or in five antesepalous fascicles, free from or adnate to petal bases. Anthers dorsifixed (outer stamens) or basifixed (inner stamens), versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective usually not protruding. Tapetum secretory? Inner stamens often with smaller anthers, or sometimes c 40 intrastaminal staminodia without anthers present; innermost staminodia often markedly shorter than remainder and with nectariferous glands at base. Staminodia fused with adjacent stamens into tube encircling gynoecium.

Pollen grains Microsporogenesis simultaneous? Pollen grains (2–)3(–6)-colporate (sometimes parasyncolpate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, finely reticulate, rugulate or verrucate.

Gynoecium Pistil composed of three or four (to six) (Caryocar) or eight to c. 15(–20) (Anthodiscus) connate carpels. Ovary superior, quadrilocular (to sexalocular) (Caryocar) or 8- to c. 15(–20)-locular (Anthodiscus), synascidiate. Stylodia three or four (Caryocar) or eight to at least twelve (Anthodiscus), free, long and filiform (Caryocar) or short (Anthodiscus); each style in Anthodiscus supported by one vascular bundle from adjacent carpels. Stigmas punctate, unicellular-papillate, type? Pistillodium absent.

Ovules Placentation axile. Ovule one per carpel, hemianatropous to weakly campylotropous, ascending, epitropous, bitegmic (Caryocar) or unitegmic (Anthodiscus), weakly crassinucellar. Micropyle usually bistomal (sometimes endostomal). Outer integument three to five cell layers thick. Inner integument five to seven cell layers thick. Integument in Anthodiscus four or five cell layers thick. Obturator absent. Endothelium absent. Megasporangium cytoplasm-rich and filled with starch grains. Apical epidermal cells of megasporangium radially elongate. Megagametophyte monosporous, Polygonum type? Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A drupe or drupaceous schizocarp (in Caryocar with radiating fibres) with single-seeded pyrenes. Pericarp carnose to more or less lignified.

Seeds Aril absent. Testa indistinct, vascularized, sometimes aerenchymatous. Exotegmen? Endotegmen? Perisperm not developed. Endosperm thin or absent. Embryo well differentiated, chlorophyll? Hypocotyl in Anthodiscus very large, oily and proteinaceous, spirally twisted. Cotyledons two, small, inflexed. Germination phanerocotylar.

Phytochemistry Lupeol, oleanolic acid derivatives, ellagic and gallic acids, and phytosterols (sitosterol, stigmasterol) present. Cyanogenic compounds not found.

Distribution Central Africa, southern India, Sri Lanka, the Andaman Islands, Southeast Asia, Malesia to New Guinea, islands in southwestern Pacific.

Vegetative anatomy Ectomycorrhiza present in Bhesa. Phellogen ab initio usually superficial? (sometimes cortical). Vessel elements with scalariform perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements ? with simple or bordered pits, non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma apotracheal diffuse-in-aggregates, or paratracheal scanty, reticulate, or banded. Sieve tube plastids S type? Nodes 5:5, pentalacunar with five leaf traces (Bhesa). Prismatic calciumoxalate crystals abundant.

Leaves Alternate (in Bhesa spiral; in Centroplacus distichous), simple, entire, in Bhesa with conduplicate ptyxis. Stipules cauline (lateral), small, in Centroplacus persistent, in Bhesa large and almost enclosing stem/branch, caducous; leaf sheath absent. Colleters present in Bhesa (absent in Centroplacus). Petiole in Bhesa with apical pulvinus. Petiole vascular bundle transection in Bhesa U-shaped or flat-annular; petiole with two or three medullary bundles and sometimes wing bundles. Venation pinnate, in Centroplacus brochidodromous; secondary veins in Bhesa stout, ascending (reticulate venation); tertiary veins in Bhesa closely scalariform. Stomata anisocytic (Centroplacus) or laterocytic (Bhesa). Cuticular wax crystalloids? Mesophyll in Centroplacus with sclerenchymatous idioblasts (containing different kinds of sclereids). Leaf margin in Centroplacus indistinctly serrate, in Bhesa entire.

Inflorescence Male inflorescences in Centroplacus axillary branched panicle, female inflorescences racemiform-subpaniculate; in Bhesa terminal? racemose simple or branched. Bracts very small.

Flowers Actinomorphic, small. Pedicels articulated. Hypogyny. Sepals five, in Centroplacus with imbricate aestivation, persistent, free. Centroplacus: petals in male flowers five, with imbricate aestivation, free, absent in female flowers; Bhesa: petals five, with contorted aestivation, free. Nectariferous disc in Centroplacus extrastaminal, in male flowers cupular, in female flowers acetabuliform, with five alternisepalous lobes; nectariferous disc in Bhesa often lobate.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals. Anthers basifixed (Centroplacus), non-versatile?, tetrasporangiate, extrorse (Bhesa) to introrse, longicidal (dehiscing by longitudinal slits, in Centroplacus two obliquely apical slits); connective in Centroplacus well developed. Tapetum secretory? Female flowers in Centroplacus with very small antesepalous staminodia (absent in Bhesa).

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate (Centroplacus) or semitectate (Bhesa), with brevicolumellate infratectum, psilate, in Centroplacus finely reticulate or perforate, in Bhesa finely striate.

Gynoecium Pistil composed of three (Centroplacus) or two (Bhesa) connate carpels. Ovary superior, trilocular (Centroplacus) or usually bilocular (rarely unilocular; Bhesa). Style in Centroplacus single, short with three diverging and somewhat recurved branches, stylodia in Bhesa two, almost entirely free. Stigmas small, capitate, slightly widened, type? Male flowers in Centroplacus with entire or trilobate hairy pistillodium; pistillodium absent in Bhesa.

Ovules Placentation subapical (Centroplacus) or basal (Bhesa). Ovules two per carpel, collateral, anatropous, erect apotropous (Bhesa) or epitropous (Centroplacus), bitegmic?, crassinucellar? Micropyle exostomal (Bhesa). Outer integument six to eight cell layers thick (Bhesa). Inner integument four or five cell layers thick (Bhesa). Endostome lignified, more or less protruding (Centroplacus). Obturator absent. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A septicidal and sometimes loculicidal capsule with single-seeded locules (one seed aborting), dehiscing from base and with persistent calyx (columella absent).

Seeds Aril exostomal-funicular, fleshy, red to orange, completely or almost completely enclosing seed (Bhesa). Carunculus narrowly elongate, red (Centroplacus). Exotesta in Centroplacus with thickened outer cell walls. Endotesta? Exotegmic cells ribbon-shaped with thick walls, in Bhesa massive. Mesotegmic cells flattened and orientated at right angles (Centroplacus). Endotegmic cells in Centroplacus more or less thick-walled. Perisperm not developed. Endosperm in Centroplacus copious, carnose. Embryo minute, short, chlorophyll? Cotyledons two. Germination phanerocotylar?

Brown in J. H. Tuckey, Narr. Exped. Zaire: 433. 5 Mar 1818 [’Chrysobalaneae’], nom. cons.

Licaniaceae Martinov, Tekhno-Bot. Slovar: 336. 3 Aug 1820 [’Licaneae’]; Chrysobalanales Link, Handbuch 2: 72. 4-11 Jul 1829 [’Chrysobalaneae’]; Hirtellaceae Horan., Char. Ess. Fam: 152. 30 Jun 1847

Fossils Fossils of Chrysobalanaceae have been found Eocene and later layers in North, Central and South America and tropical Asia.

Habit Usually bisexual (rarely andromonoecious or gynomonoecious), evergreen trees or shrubs. Lenticels abundant.

Vegetative anatomy Phellogen ab initio usually superficially (sometimes deeply) seated. Primary medullary strands narrow (entirely or largely uniseriate). Vessel elements with simple perforation plates; lateral pits alternate, usually with simple pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate, usually heterocellular (rarely homocellular). Axial parenchyma apotracheal, diffuse or diffuse-in-aggregates, or paratracheal scanty, reticulate, or banded. Tyloses abundant. Sieve tube plastids S type. Nodes 5:5, pentalacunar with five leaf traces. Wood usually with silica bodies and grains. Prismatic calciumoxalate crystals sometimes abundant.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, stellate or arachnoid; stalked and unstalked glands (also peltate-lepidote) present on calyx and leaves.

Leaves Alternate (spiral or distichous), simple, entire, with conduplicate (sometimes flat-conduplicate) ptyxis. Stipules often petiolar or intrapetiolar; leaf sheath absent. Petiole often pulvinate at one or both ends. Petiole vascular bundle transection annular; petiole often with medullary plates and wing bundles; petiolar anatomy often complex. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids? Lamina often with flattened abaxial glands (extrafloral nectaries), especially near base. Myrmecodomatia (pouches formed when lamina rolls over onto itself creating two spherical spaces at base) present in some species of Hirtella; myrmecodomatia associated with enlarged extrafloral nectaries on stipules and bracts, longer stomatal apertures, enlarged parenchymatous and epidermal cells, and more numerous lignified sclerenchyma fibres. Epidermis with or without silica bodies, with or without mucilaginous idioblasts. Mesophyll sometimes with sclerenchymatous idioblasts. Leaf margin usually entire (rarely serrate). Extrafloral nectaries sometimes present on stipules, petiole and/or lamina.

Inflorescence Terminal or axillary, cymose or racemose of various shape (simple or compound raceme etc.; flowers rarely solitary). Extrafloral nectaries sometimes present on bracts or pedicels.

Flowers Actinomorphic to obliquely zygomorphic, usually small. Pedicel articulated. Hypogyny. Sepals five, with imbricate quincuncial aestivation, connate in lower part into tubular hypanthium-like structure (“floral cup”); median sepal adaxial. Petals one to five, usually with imbricate quincuncial (sometimes cochlear) aestivation, shortly clawed, free, adnate at margin of “hypanthium” (petals rarely absent). Nectariferous disc annular or semicircular, intrastaminal, inserted inside or along apex of “hypanthium”.

Androecium Stamens (two to) five to c. 300, usually long exserted, in zygomorphic flowers concentrated to one side of flower (lateral antesepalous stamens often larger than remainder); abaxial stamens most developed. Filaments inflexed in bud, free or more or less connate all together or in three to 20 staminal fascicles, free from tepals, inserted at margin of hypanthium-like structure. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often dorsally thickened? Tapetum secretory. Intrastaminal staminodia often present.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–4)-colp(or)ate, shed as monads, bicellular at dispersal. Exine with very little patterning on walls, with ? infratectum, usually scabrate to verrucate.

Gynoecium Pistil composed of one to three carpels, fused only by common lateral to gynobasic style; usually only abaxial carpel developed leading to pseudomonomery (remaining carpels usually degenerated), often inserted on one side of “hypanthium”. Ovary superior, usually unilocular; locule sometimes divided by secondary septa. Style single, simple, lateral to gynobasic. Stigma usually simple, usually punctate (rarely trilobate), papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation basal-axile. Ovules two per carpel, anatropous, ascending, epitropous (antitropous), collateral or distributed over carpellary surface, bitegmic, tenuinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument five to twelve cell layers thick. Inner integument five to twelve cell layers thick. Obturator seemingly basal (pollen tube transferring tissue perhaps functioning as obturator) or absent. Archespore multicellular. Megagametophyte consisting of an egg cell, two synergids and a central cell. Antipodal cells absent (early degenerating). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A single-seeded, usually juicy (rarely dry) drupe with persistent calyx. Endocarp often hairy inside.

Seeds Aril absent. Seeds sometimes ruminate. Testa usually well developed, vascularized (sometimes indistinct or mesotestal, sometimes multiplicative). Exotesta fibrous, collapsed (sometimes with tannins). Tegmen multiplicative. Perisperm not developed. Endosperm absent. Embryo large, well differentiated, chlorophyll? Cotyledons two, sometimes thick. Germination cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), trihydroxyflavonoids, cyanidin, delphinidin, cucurbitacins, and tannins present. Special unsaturated fatty acids present in seeds. Ellagic acid, alkaloids, saponins, and cyanogenic compounds not found.

Use Fruits (Chrysobalanus icaco, Neocarya, Parinari), seed oil, timber.

Lindley, Intr. Nat. Syst. Bot., ed. 2: 74. 13 Jun 1836 [’Guttiferae, vel Clusiaceae’], nom. cons.

Guttiferae Juss., Gen. Plant: 255. 4 Aug 1789, nom. cons. et nom. alt.; Garciniaceae Bartl., Ord. Nat. Plant.: 222, 292. Sep 1830 [’Garcinieae’]; Cambogiaceae Horan., Prim. Lin. Syst. Nat.: 98. 2 Nov 1834 [’Cambogiaceae (Guttiferae)’]; Garciniales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 60. Sep-Oct 1835 [‘Garcinieae’]

Fossils Paleoclusia chevalieri, pentamerous flowers from the Turonian of New Jersey, has been attributed to Clusiaceae and one of the few known Cretaceous representatives of Malpighiales. The stamens (or staminodia) are grouped into five fascicles and a resin-like amorphous substance is present in (and was perhaps secreted by) the anthers. The ovary is quinquelocular and the stigma quinquelobate. The seeds are arillate. ’Pachydermites diederexii, fossil pollen of Symphonia, has been used for stratigraphic dating by the oil industry.’ Symphonioxylon, fossil wood from the Cretaceous and the Miocene in northeastern Africa and India, may be ascribed to Clusiaceae or Calophyllaceae.

Habit Usually bisexual (rarely polygamomonoecious, in e.g. Clusia and Garcinia also dioecious), evergreen trees or shrubs (in Clusia sometimes lianas or epiphytes, also with CAM physiology).

Vegetative anatomy Phellogen ab initio in roots (sub)epidermal or deeply seated; in stem superficial. Secondary lateral growth usually normal. Vessel elements usually with simple (sometimes also scalariform) perforation plates; lateral pits alternate, usually simple pits. Vestured pits often present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres usually with simple (rarely bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal or paratracheal. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Schizogenous secretory canals or cavities (and/)or glands with resin, balsam or yellow to red secretions frequent (also in cortex and medulla). Colleters numerous. Wood ray cells sometimes with silica. Crystals?

Leaves Usually opposite (sometimes alternate; rarely whorled), simple, entire, often coriaceous, with conduplicate or flat ptyxis. Stipules and leaf sheath absent. Paired modified colleters, “stipular glands”, may occur. Petiole bundle transection annular. Venation pinnate, usually eucamptodromous or brochidodromous (rarely acrodromous). Stomata paracytic. Cuticular wax crystalloids as rosettes. Lamina usually with schizogenous secretory glands or canals with resin, balsam or yellowish to reddish secretions. Leaf margin entire.

Inflorescence Terminal or axillary, cymose, often thyrsoid (flowers sometimes solitary).

Flowers Actinomorphic, often large. Hypogyny. Sepals (two to) four or five (to 20), usually with imbricate quincuncial or decussate (rarely valvate) aestivation, usually free (rarely connate at base). Petals (three or) four or five (to eight), with contorted or decussate aestivation, free (sometimes absent). Nectariferous disc usually absent (sometimes as separate units at staminal bases; in Symphonia extrastaminal nectariferous disc possibly representing antesepalous staminal whorl). Resins often frequently secreted (in, i.a., Clusia).

Androecium Stamens (four to) c. 20 to more than 100, often in distinct fascicles. Filaments stout, free, or more or less connate into five alternisepalous, antepetalous fascicles, free from tepals. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, usually introrse (sometimes extrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by pores); connective usually without glands (sometimes with small glands). Tapetum secretory. Staminodia two to more than 50, extrastaminal (sometimes producing viscid triterpenoid resin), or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes triporate or with several apertures), usually shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate? infratectum, reticulate, rugulate, fossulate, foveolate, scabrate, spinulate or psilate.

Gynoecium Pistil composed of two to five (to more than twelve) connate, often antesepalous carpels (rarely monocarpellate?). Ovary superior, bilocular to quinquelocular (to more than duodecemlocular in Garcinieae, with single-seeded locules). Stylodia two to five (to more than twelve), free, usually short (shorter than ovary), or style single, simple, or absent. Stigmas usually several (sometimes one entire stigma), usually widened (sometimes punctate) and non-papillate (rarely papillate), Wet type (exposed stigmatic area absent in Symphonieae). Pistillodium? Pollen grains in, e.g., Symphonia collected in droplet secreted through pore at apex of stylar branches.

Ovules Placentation usually axile (sometimes apical or basal; in Allanblackia parietal). Ovules (one or) two to numerous per carpel, usually anatropous (sometimes amphitropous or hemianatropous), ascending to horizontal, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad. Agamospermy present in Clusia and Garcinia mangostana.

Fruit Usually a drupe or berry (in Garcinieae and Symphonieae many-seeded; in some species of Garcinia drupe; sometimes a septicidal capsule).

Seeds Aril sometimes present. Seed coat testal or exotegmic. Testa sometimes winged; testa with only epidermis and exotegmen usually with lignified sinuous cell walls, or testa multi-layered, vascularized and complex and exotegmen usually absent. Endotegmen? Perisperm not developed. Endosperm sparse or absent. Embryo large to small, straight and fusiform or curved, rudimentary or well differentiated, with or without chlorophyll. Cotyledons two, very small or rudimentary. Hypocotylar region very enlarged, forming tigellus. Germination phanerocotylar or cryptocotylar. Radicula in species with large seeds often ephemeral and replaced by adventitious roots.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, biflavonoids, flavonoid sulphates, cyanidin, dammaranes, oleanolic acid derivatives, ellagic and gallic acids, proanthocyanidins (prodelphinidins), alkaloids, cyanogenic compounds, isoprenylated and other xanthones (euxanthone, macluraxanthone, norathyriol), polyacetate-derived anthraquinones, polyisoprenylated benzophenones and fatty acids (as resins), and syringaresinol present.

Use Ornamental plants, fruits (Garcinia mangostana, Moronobea coccinea, Platonia insignis), medicinal plants, gums and resins, seed oils, timber.

Fossils Fossil pollen assigned to Ctenolophon has been found in South America, India and Malaysia, and from Maastrichtian layers in Africa (Muller 1981, van der Ham 1989), although these records are not cited by Friis & al. (2011).

Vegetative anatomy Phellogen? Vessel elements with scalariform or reticulate perforation plates; lateral pits alternate, scalariform or opposite, bordered pits. Imperforate tracheary xylem elements ? with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal aliform, winged-aliform, confluent, vasicentric, or unilateral. Sieve tube plastids ? type. Nodes 3:3, trilacunar with three leaf traces. Cristarque cells present. Prismatic calciumoxalate crystals frequent.

Trichomes Hairs multicellular?, simple, fasciculate or stellate (stellate hairs present on leaves, stipules and tepals).

Leaves Opposite, simple, entire, coriaceous, with ? ptyxis. Stipules interpetiolar, early caducous, without colleters; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate (arcuate and anastomosing). Stomata anomocytic or anisocytic. Cuticular waxes absent. Leaf margin entire.

Flowers Actinomorphic. Pedicel articulated. Hypogyny. Sepals five, with imbricate quincuncial aestivation, persistent, connate at base. Petals five, with contorted aestivation, fleshy, spoon-shaped at base, caducous, free. Nectariferous disc extrastaminal, cupular, with stomata.

Androecium Stamens 5+5, diplostemonous. Filaments of two different lengths, inserted in lower part of adaxial side of nectariferous disc, basally fused into tube, adnate at base to petals (epipetalous). Ten lobes alternating with stamens and forming corona-like tube at base on dorsal side of stamens. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective wide and thick, protruded. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3–9-stephanocolpate or 3–9-stephanocolpor(oid)ate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, perforate, spinulate. Pollen grains of unique type, almost square in polar view and with extremely wide colpi.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular, synascidiate. Style single, bifid. Stigmas capitate, unicellular-papillate, type? Pistillodium absent.

Ovules Placentation lateral, apical, axile. Ovules two per carpel, anatropous, pendulous, epitropous, bitegmic, (weakly) crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Integuments lobate. Outer integument approx. five cell layers thick. Inner integument ten or eleven cell layers thick. Obturator placental. Endothelium present. Megasporangium disintegrating. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A single-seeded nut (nut-like capsule?) with lignified pericarp and persistent, accrescent, swollen calyx. Seed persistent on thin central funicular columella.

Seeds Arillode hairy, enclosing lower half of seed. Exotestal cells palisade, with thickened outer wall. Endotesta? Exotegmen fibrous. Endotegmen? Perisperm not developed. Endosperm copious. Embryo straight, well differentiated, chlorophyll? Cotyledons two, very large, plicate. Germination?

Baillon in von Martius, Fl. Bras. 12(1): 365. 1 Apr 1886 [‘Dichapetaleae’], nom. cons.

Chailletiaceae R. Br. in J. H. Tuckey, Narr. Exped. Zaire: 442, 443. 5 Mar 1818 [’Chailleteae’]; Chailletiales Link, Handbuch 2: 123. 4-11 Jul 1829 [‘Chailletiaceae’]

Habit Usually at least morphologically bisexual (at least sometimes functionally monoecious or dioecious), evergreen trees, shrubs or lianas. Sometimes xerophytic. Lenticels often numerous.

Vegetative anatomy Phellogen ab initio superficial. Pericyclic envelope interrupted. Vessel elements with simple and/or scalariform perforation plates; lateral pits usually alternate (in Tapura often scalariform or intermediary), bordered pits. Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually paratracheal aliform, lozenge-aliform, winged-aliform, vasicentric, or confluent (sometimes apotracheal diffuse). Sieve tube plastids S type?; sieve tubes with non-dispersive protein bodies. Nodes? Pericyclic envelope interrupted. Resinous substances produced? Prismatic calciumoxalate crystals abundant.

Trichomes Hairs unicellular, simple, verrucose-papillate; glandular hairs (glands) sometimes present.

Leaves Alternate (spiral), simple, entire, sometimes coriaceous, with ? ptyxis. Stipules intrapetiolar, often fimbriate, usually caducous; leaf sheath absent. Petiole articulated. Petiole vascular bundle transection arcuate. Venation pinnate, brochidodromous; secondary veins strongly curved. Stomata usually paracytic (sometimes anomocytic). Cuticular wax crystalloids? Domatia as pits. Epidermis with mucilaginous idioblasts. Mesophyll with or without mucilaginous idioblasts. Lamina sometimes with abaxial flattened glands and sometimes with mucilage cells. Leaf margin entire. Extrafloral nectaries often present on adaxial or abaxial side of lamina in Dichapetalum.

Inflorescence Axillary (to petiolar; in two species of Dichapetalum and one species of Tapura epiphyllous, arising from petiole), cymose (often fasciculate).

Flowers Usually actinomorphic (in Tapura obliquely zygomorphic), small. Pedicel often articulated. Hypanthium-like structure (“floral cup”) present or absent. Usually hypogyny to half epigyny (rarely epigyny). Sepals four or five, with imbricate quincuncial aestivation, usually free (sometimes connate at base). Petals four or five, with imbricate, involute or valvate to somewhat contorted (rarely open) aestivation, usually deeply bifid, sometimes clawed, blackening when dry, usually free (rarely more or less connate). Nectariferous disc annular when petals connate or as four or five antepetalous scale-like nectariferous glands between staminodia, inserted at petal bases (nectariferous disc in Tapura lobate, semicircular).

Androecium Stamens usually four or five (in Tapura three), haplostemonous, antesepalous, alternipetalous (in Tapura three fertile stamens and two staminodia). Filaments free or connate at base into tube, free from or adnate to tepals. Anthers dorsifixed to almost basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often dorsally thickened. Tapetum secretory. Female flowers with staminodia; staminodia (intrastaminal?) in male flowers of Tapura two or five, antesepalous.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular? at dispersal. Exine semitectate, with columellate infratectum, reticulate (sometimes microreticulate) to retipilate.

Gynoecium Pistil composed of (two or) three (or four) connate carpels; carpel synascidiate. Ovary usually superior to semi-inferior (rarely inferior), (bilocular or) trilocular (or quadrilocular), with central columella and sometimes secondary septa. Style usually single, simple or branched (stylodia rarely two to four, free). Stigma capitate or trilobate, often with recurved lobes, papillate, Wet type. Male flowers with pistillodium.

Ovules Placentation axile to apical. Ovules two per carpel, anatropous, pendulous, epitropous (antitropous, with micropyle above), bitegmic, tenuinucellar. Micropyle bistomal, Z-shaped (zig-zag), or endostomal. Outer integument three to five cell layers thick. Inner integument six to eight cell layers thick. Obturator apical, with long multicellular papillae or hairs, or funicular. Hypostase absent. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit An often flattened usually unilocular (rarely bilocular or trilocular and lobed) dry or sometimes juicy single-seeded drupe (sometimes a capsule) with dense, short, erect and often golden yellow hairs and persistent calyx.

Seeds Aril or carunculus present or absent. Testa multiplicative, vascularized, only consisting of enlarged tanniniferous, sometimes divided, exotestal cells and remnants of vascular bundles; testal cells often with stellate calcium oxalate crystals. Endotesta? Exotegmen absent. Endotegmen? Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, oily, with chlorophyll. Cotyledons two. Germination cryptocotylar.

Phytochemistry Insufficiently known. Pyridine alkaloids sometimes present. Some genera (i.a. Dichapetalum) with strongly toxic fluoroacetic acid (fluoroacetate). Saponins not found.

Cryptaceae Raf. in Ann. Gén. Sci. Phys. Bruxelles 5: 349. Jul-Sep 1820 [‘Cryptinia’]; Elatinales Cambess. in C. F. P. von Martius, Consp. Regn. Veg.: 54. Sep-Oct 1835 [‘Elatineae’]; Alsinastraceae Rupr., Fl. Ingr. 1: 194. Mai 1860 [’Alsinastreae’], nom. illeg.

Distribution Cosmopolitan except polar areas, with their largest diversity in tropical regions.

Fossils Curved seeds with reticulate surface, resembling those in Elatine, are known from the Pliocene of Europe.

Habit Bisexual, usually perennial or annual herbs (Bergia suffruticosa is suffrutescent). A large number of species are aquatic, whereas others are amphibious helophytes.

Vegetative anatomy Phellogen ab initio usually superficial (in Bergia inner-cortical). Endodermis prominent in species of Elatine. Secondary lateral growth normal or absent. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits? Imperforate tracheary xylem elements tracheids with bordered pits, non-septate? (also vasicentric tracheids). Wood rays uniseriate to multiseriate, heterocellular? Axial parenchyma apotracheal diffuse. Sieve tube plastids S₀ type, without starch or protein inclusions. Nodes 1:1, unilacunar with one leaf trace. Mucilage cells and resin-producing cells present. Secretory cells with tanniniferous substances (resinous latex) present in Bergia. Druses and solitary crystals present.

Trichomes Hairs usually absent (when present unicellular or uniseriate, simple, basifixed and sometimes gland-tipped); glandular hairs abundant in Bergia

Leaves Usually opposite (in Elatine alsinastrum verticillate), simple, entire, with ? ptyxis. Stipules interpetiolar, minute, membranous; leaf sheath absent. Colleters present. Petiole vascular bundle simple. Venation pinnate. Stomata usually paracytic (rarely anomocytic or tetracytic), with four to eight irregularly formed subsidiary cells. Cuticular wax crystalloids? Epidermis with or without mucilaginous idioblasts. Lamina with or without secretory cavities. Leaf margin serrate, crenate or entire, often with multicellular glands along margin.

Flowers Actinomorphic, small. Hypogyny. Sepals (two or) three to five (or six), with imbricate aestivation, free or connate at base; median sepal (when three sepals) abaxial. Petals (two or) three to five (or six), with contorted or imbricate aestivation, persistent, free. Nectary absent. Disc absent.

Androecium Stamens (two or) three to five (or six), haplostemonous, antesepalous, or (four to) six to ten (to twelve), in one or two whorls, diplostemonous, with outer stamens antesepalous and inner stamens alternisepalous (inner staminal whorl sometimes absent). Filaments free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate, shed as monads, bicellular (Bergia) or tricellular (Elatine) at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of two to five (or six) connate antesepalous carpels; median carpel (when three carpels) abaxial. Ovary superior, bilocular to quinquelocular (or sexalocular; septa sometimes not reaching apex). Stylodia two to five (or six), free. Stigmas capitate, papillate, type? Pistillodium absent.

Ovules Placentation apical, pendulous. Ovules (two to) numerous per carpel, anatropous, epitropous, ascending or horizontal, bitegmic, weakly crassinucellar. Micropyle bistomal, sometimes Z-shaped (zig-zag). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Archespore multicellular. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad.

Seeds Aril absent. Operculum present. Seed coat exotegmic. Testa collapsed? Exotegmic cells with sinuous anticlinal walls, lignified. Endotegmen? Perisperm not developed. Endosperm thin (Bergia) or absent (Elatine). Embryo straight or somewhat curved, large, fusiform, well differentiated, chlorophyll? Cotyledons two, short. Germination phanerocotylar.

Phytochemistry Flavonols, ellagic acid, tannins, and proanthocyanidins (prodelphinidins) present. Alkaloids and saponins not found.

Elatine alsinastrum is sister to the remaining species of Elatine, based on morphological characters (Razifard & al. 2017).

Kunth in von Humboldt, Bonpland et Kunth, Nov. Gen. Sp. Plant. 5, ed. 4^o: 175; ed. fol.: 135. 25 Feb 1822 [‘Erythroxyleae’], nom. cons.

Erythroxylales Link, Handbuch 2: 339. 4-11 Jul 1829 [‘Erythroxyleae’]; Nectaropetalaceae (H. Winkl.) Exell et Mendonça in Bol. Soc. Brot., ser. 2, 25: 105. 1951

Habit Usually bisexual (rarely dioecious), evergreen small trees or shrubs. Branches often covered with distichous scale-like rudimentary leaves. Buds perulate.

Vegetative anatomy Mycorrhiza absent. Phellogen ab initio superficial. Young stems and branches with cortical vascular bundles. Primary vascular tissue cylinder, without separate bundles. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Vestured pits absent (Erythroxylum). Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, winged-aliform, vasicentric, or confluent. Secondary phloem often stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids Pc (PV) type, with numerous square or polygonal protein crystalloids. Nodes 3:3, trilacunar with three leaf traces, with lateral vascular bundles originating relatively long before central bundle and forming cortical bundles. Sclereids present. Wood usually with silica grains. Parenchyma and bundle envelopes usually with cristarque cells (sometimes absent). Prismatic calciumoxalate crystals frequent.

Leaves Usually alternate (spiral or distichous; in Aneulophus opposite), simple, entire, with involute ptyxis. Stipules usually at least partially interpetiolar (in Exythroxylum intrapetiolar), sheathing, in all species with basal adaxial gum-secreting colleters; leaf sheath absent. Petiole vascular bundle transection arcuate or annular; petiole with medullary and adaxial bundles. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type). Epidermis with or without mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts. Leaf margin entire.

Flowers Usually actinomorphic (in Aneulophus zygomorphic?), often small. Pedicel usually articulated. Hypanthium present in Nectaropetalum. Hypogyny. Sepals five, with imbricate quincuncial or valvate aestivation, persistent, connate at base. Petals five, with imbricate or contorted aestivation, caducous, usually with adaxial fimbriate bilobate ligule at base, free or connate at base. Nectariferous glands present on abaxial side of staminal tube. Disc absent. Heterostyly frequent.

Androecium Stamens 5+5, usually obdiplostemonous (sometimes diplostemonous). Filaments connate into tube around pistil at least at base, free from tepals, often with two different lengths. Anthers dorsifixed to basifixed, versatile?, tetrasporangiate, latrorse (or introrse?), longicidal (dehiscing by longitudinal slits); connective often thickened. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tricolpate), shed as monads, tricellular at dispersal. Exine semitectate, with columellate infratectum, finely reticulate to punctate.

Gynoecium Pistil composed of (two or) three (or four) connate carpels; usually only adaxial carpel (rarely all carpels) fertile. Ovary superior, usually with only one fertile locule, synascidiate. Stylodia (two or) three (or four), free or more or less connate, sometimes with canal. Stigmas capitate, unicellular-papillate, Dry type. Pistillodium absent.

Ovules Placentation apical to axile. Ovule one (or two) per carpel, anatropous to hemianatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument two to five cell layers thick. Inner integument five to nine cell layers thick. Hypostase absent. Parietal tissue two to four cell layers thick. Endothelium usually present. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad.

Fruit Usually a single-seeded drupe (in Aneulophus a septicidal capsule) with persistent calyx and stamens.

Seeds Arilloid (carunculus?), formed from exostome, present in Aneulophus. Seed coat exotegmic. Exotestal cells thickened, elongate. Endotesta crushed. Tegmen often multiplicative (in Aneulophus thin). Exotegmen fibrous. Endotegmen as pigmented endothelium. Perisperm not developed. Endosperm very copious, starchy (rarely absent). Embryo straight, well differentiated, with chlorophyll. Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, ethereal oils (in wood), non-hydrolyzable tannins, chlorogenic acid, tropane (hygrolinic) alkaloids (tropane-3α and tropane-3β-ols, tropacocaine, scopolamine oxides, hydroxytropines, teloidines, ecgonines, norecgonines, etc.), pyrrolizidine alkaloids, and saponins present. Ellagic acid and cyanogenic compounds not found.

Use Medicinal plants, cocaine (Erythroxylum coca, E. novogranatense), timber, tar, dyeing substances.

Tithymalaceae Vent., Tabl. Règne Vég. 3: 483. 5 Mai 1799 [’Tithymaloideae’]; Euphorbiales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 237. Jan-Apr 1820 [‘Euphorbiaceae’]; Mercurialaceae Bercht. et J. Presl, Přir. Rostlin: 237. Jan-Apr 1820 [’Mercurialideae’]; Ricinaceae Martinov, Tekhno-Bot. Slovar: 547. 3 Aug 1820 [’Ricini’]; Trewiaceae Lindl., Intr. Nat. Syst. Bot., ed. 2: 174. 13 Jun 1836; Tragiaceae Raf., Fl. Tellur. 4: 111. med 1838 [’Tragides’]; Acalyphaceae Juss. ex Menge., Cat. Plant. Grudent. Gedan.: 172. 1839 [’Acalyphinae’]; Crotonopsida Brong., Enum. Pl. Mus. Paris: xxiii, 79. 12 Aug 1843 [‘Crotonineae’]; Bertyaceae J. Agardh, Theoria Syst. Plant.: 190. Apr-Sep 1858; Crotonaceae J. Agardh, Theoria Syst. Plant.: 258. Apr-Sep 1858 [’Crotoneae’]; Hippomanaceae J. Agardh, Theoria Syst. Plant.: 244. Apr-Sep 1858 [’Hippomaneae’]; Ricinocarpaceae (Müll. Arg.) Hurus. in J. Fac. Sci. Univ. Tokyo, ser. III, 6: 224. 15 Aug 1954; Euphorbianae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992; Cheilosaceae Doweld, Tent. Syst. Plant. Vasc.: xxxi. 23 Dec 2001

Distribution Cosmopolitan except polar areas, although mainly tropical, with their largest species diversity (Euphorbia) in southern Africa, the Mediterranean and the irano-turanian regions, and southern North America.

Fossils Fossil fruits and seeds of Euphorbiaceae from the Eocene of England (the London Clay and the Pipe-Clay in Dorset) have been described (e.g. as Euphorbiospermum and Euphorbiotheca). From Oligocene and younger layers in Australia there are likewise fossils assigned to Euphorbiaceae. Crepetocarpon from the Eocene of North America may belong in Euphorbiaceae-Euphorbioideae. Fossil wood, probably of euphorbiacean origin, is frequently documented from Palaeogene and Neogene layers on several continents (also New Zealand).

Habit Monoecious or dioecious, evergreen or deciduous trees or shrubs, perennial or annual herbs (rarely lianas; many species are stem succulents and xerophytes). CAM and C₄ physiologies present in many species (i.a. Chamaesyce subclade of Euphorbia).

Vegetative anatomy Phellogen ab initio outer-cortical or pericyclic. Primary vascular tissue bicollateral or centrifugal. Cortical and medullary vascular bundles sometimes present. Secondary lateral growth usually normal (sometimes anomalous, from cylindrical cambium) or absent. Vessel elements often in multiples. Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate, simple pits. Vestured pits sometimes present. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate (in Bernardia also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, eller paratracheal scanty, reticulate, scalariform, vasicentric, or banded, or absent. Tyloses sometimes frequent. Sieve tube plastids S type; sieve tubes with non-dispersive protein bodies. Nodes usually 3:3, trilacunar with three leaf traces (rarely 1:1, unilacunar with one trace, or ≥5:≥5, multilacunar with five or more traces). Articulated or inarticulated laticifers with gums (rubber; usually absent in Acalyphoideae). Secretory canals with tannins. Silica bodies present in many species. Prismatic crystals often abundant (also in axial parenchyma and/or wood ray cells); acicular crystals, druses, styloids, crystal sand or other types of calciumoxalate crystals present or absent.

Trichomes Hairs unicellular or multicellular, malpighiaceous hairs (unicellular T-shaped; Argythamnia, Chiropetalum, Rhodothyrsus), simple, furcate, stellate, candelabra-shaped, dendritic, peltate or lepidote; glandular hairs often present; stinging hairs sometimes present (in, e.g., Cnidoscolus and Tragia).

Leaves Usually alternate (spiral or distichous) or opposite (rarely verticillate), usually simple (sometimes palmately compound), entire or lobed, with various ptyxis (sometimes absent). Stipules usually cauline (sometimes hair-like or modified into glands; in many species of Euphorbia modified into spines; rarely absent); leaf sheath absent. Petiole often with distal pulvinus, sometimes with paired or unpaired extrafloral nectaries. Petiole vascular bundle transection arcuate, annular etc. Venation usually palmate (sometimes pinnate). Stomata usually paracytic (sometimes anisocytic, parallelocytic or anomocytic). Cuticular wax crystalloids as rosettes of platelets (Fabales type), or cuticular wax as crust (especially in succulents). Epidermis with or without mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts. Styloids present in some genera. Secretory cavities absent. Leaf margin entire or serrate with simple veins proceeding into persistent transparent leaf teeth. Subbasal glands present or absent; extrafloral nectaries sometimes present on lamina.

Inflorescence Terminal or axillary, cymose of various shapes (panicle, thyrsoid, fascicle, corymb, raceme-, catkin- or spike-like, etc.) consisting of monochasia or dichasia. Bracts sometimes involucral, sometimes large and showy, occasionally resin-producing (e.g. Dalechampia). Inflorescence in Euphorbia pseudanthial cyathium (thyrsoid), each with one terminal female partial inflorescence with one female flower consisting of one tricarpellate pistil, surrounded by four or five cincinni of male flowers, each consisting of one stamen; five usually nectariferous bracts inserted on abaxial side of each cyathium. Female flowers in Ricinus in distal part of inflorescence.

Flowers Actinomorphic, small. Hypogyny. Sepals (two or) three to six (to twelve), with valvate or imbricate aestivation, usually free (sometimes connate at base; absent in some genera; female flowers in Excoecaria with three tepals). Petals (two or) three to six (to eight), with valvate or imbricate aestivation, usually free, or absent. Nectariferous disc intrastaminal or extrastaminal, annular or subdivided into separate glands, or absent.

Androecium Stamens one to more than 50 (to more than 1.000). Filaments usually free (sometimes connate; in Ricinus fused into branched fascicles), free from tepals, in some genera apically split and branched. Anthers basifixed to dorsifixed, often versatile, usually tetrasporangiate (rarely disporangiate), extrorse or introrse, longicidal (dehiscing by longitudinal slits). Tapetum usually secretory (sometimes amoeboid-periplasmodial). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes colpate, porate or inaperturate), shed as monads, bicellular or tricellular at dispersal. Exine tectate, semitectate or intectate, with columellate infratectum, perforate or reticulate, echinate, verruculate, spinulate or smooth.

Gynoecium Pistil composed of (two or) three (to numerous) connate carpels; median carpel usually abaxial. Ovary superior, (bilocular or) trilocular (to multilocular). Stylodia (two or) three (to numerous), free or connate at base, simple or branched. Stigmas relatively large, often branched or with adaxial furrow, papillate or non-papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation apical. Ovules one (or two) per carpel, usually anatropous (sometimes hemianatropous or amphitropous), pendulous, usually epitropous (sometimes apotropous), bitegmic, crassinucellar. Micropyle usually exostomal (sometimes bistomal). Outer integument three to many cell layers thick. Inner integument (three or) four to 25 cell layers thick. Nucellar cap present; megasporangium usually with nucellar beak protruding through micropyle and reaching obturator. Placental obturator present between stylar canal and micropyle forming roof above micropyle. Hypostase present. Megagametophyte monosporous, disporous or tetrasporous, Polygonum type, Acalypha type or other types (Allium type?, Drusa type?, Fritillaria type?, Penaea type?). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes proliferating (in at least Jatropha sometimes up to five cells). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually onagrad (sometimes solanad? or piperad?).

Fruit Usually a septicidal (sometimes loculicidal) capsule (often explosively dehiscent) or usually tripartite (rarely bipartite or multipartite) schizocarp (with carpels, cocci, sometimes elastically dehiscing from central columella; mesocarps often separating from endocarp; rarely a berry or drupe).

Seeds Seeds often large, sometimes pachychalazal. Micropylar carunculus usually present. Aril present in some species. Seed coat usually exotegmic (sometimes endotestal). Exotesta usually palisade. Endotesta cells with amorphous calciumcarbonate. Exotegmen often palisade, consisting of malpighian-like lignified cells (Crotonoideae). Endotegmen absent. Perisperm not developed. Endosperm usually copious, often oily (sometimes absent). Embryo straight or curved, usually well differentiated, with or without chlorophyll. Cotyledons two, usually longer and wider than radicula. Germination phanerocotylar or cryptocotylar.

DNA Plastid gene infA lost/defunct (Hevea). Mitochondrial coxI intron present.

Phytochemistry Flavonols (kaempferol, quercetin), triterpenes (cucurbitacins), cocarcinogenic phorbol esterditerpenes, ethereal oils and resins, ellagic and gallic acids, ellagitannins (geraniin, mallotussic acid), p-coumaric acid, caffeic acid, alkaloids (e.g. ricinine, in Croton benzylisoquinoline alkaloids), cyanogenic glycosides derived from nicotinic acid or valine/isoleucine, polyacetate-derived arthroquinones (Clutia), ferulic acid, and lectins (hemagglutinins) present. Proanthocyanidins not found. Saponins? Aluminium accumulated in some species.

Use Ornamental plants, starch sources (Manihot), medicinal plants, rubber (Hevea), seed oils (Ricinus), timber, poisons.

Systematics Euphorbiaceae are sister to Rafflesiaceae.

The phylogenetic relationships among the genera of Acalyphoideae, Crotonoideae and Euphorbioideae are still very insufficiently known.

Cheilosoideae (Müll.-Arg.) K. Wurdack et Petra Hoffm. in Amer. J. Bot. 92: 1413. 27 Jul 2005

2/7. Cheilosa (1; C. montana; West Malesia), Neoscortechinia (6; N. angustifolia, N. forbesii, N. kingii, N. nicobarica, N. philippinensis, N. sumatrensis; southern Burma, the Nicobar Islands, Malesia to New Guinea, the Bismarck Archipelago and Solomon Islands). – Southern Burma, the Nicobar Islands, Malesia to New Guinea, Solomon Islands. Stamens five to twelve. Pollen grains echinate. Pistil composed of sometimes two connate carpels. Outer integument eight to ten cell layers thick. Inner integument eight to twelve cell layers thick. Carunculus absent. Testa with vascular bundles. Endosperm present.

[Suregada clade+[Adenoclineae+[Acalyphoideae+[Crotonoideae+Euphorbioideae]]]]

Suregada clade

1–2/c 30. Suregada (c 30; tropical Africa, Madagascar, tropical Asia to tropical Australia and islands in the Pacific), Cladogelonium (1; C. madagascariense; Madagascar; in Suregada?). – Tropical regions in the Old World. Pollen grains pantoporate, acolumellate.

[Adenoclineae+[Acalyphoideae+[Crotonoideae+Euphorbioideae]]]

Adenoclineae (Müll.-Arg.) G. L. Webster in Taxon 24: 598. 19 Dec 1975

6/67. Endospermum (11; Southeast Asia, Malesia to Fiji); Klaineanthus (1; K. gaboniae; Nigeria to Gabon), Tetrorchidium (23; tropical Africa, Mexico, Central America, Jamaica, tropical South America), Adenocline (8; southern Africa north to Malawi), Ditta (2; D. maestrensis, D. myricoides; Cuba, Hispaniola, Puerto Rico); Omphalea (22; tropical regions on both hemispheres). – Pantropical, southern Africa. Endospermum may be sister to the remaining Adenoclineae.

[Acalyphoideae+[Crotonoideae+Euphorbioideae]]

Outer integument six to approx. ten cell layers thick. Phorbol esters (phorbol diterpenes) sometimes present.

Acalyphoideae (Kunth) Beilschm. in Flora 16(Beibl. 7): 61, 104. 14 Jun 1833 [‘Acalypheae’]

c 100/2.000–>2.035. Acalypheae Dumort., Anal. Fam. Plant.: 45. 1829. Acalypha (450–455; tropical and subtropical regions on both hemispheres); Claoxylon (c 115; Madagascar, tropical Asia, northern and eastern Australia, Melanesia, the Hawaiian Islands), Crotonogynopsis (2; C. akeassii, C. usambarica; tropical Africa), Discoclaoxylon (4; D. hexandrum, D. occidentale, D. pedicellare, D. pubescens; one species in western tropical Africa, three species endemic to São Tomé); Erythrococca (c 40; tropical and southern Africa, southern Arabian Peninsula), Micrococca (12; tropical Africa, Madagascar, the Arabian Peninsula, tropical Asia to the Malay Peninsula); Cleidion (30–35; tropical regions on both hemispheres), Sampantaea (1; S. amentiflora; Thailand, Cambodia), Wetria (2; W. insignis: southern Burma, Thailand, West Malesia, the Philippines; W. australiensis: Papua New Guinea, northeastern Queensland); Dysopsis (3; D. glechomoides, D. hirsuta, D. paucidentata; the Andes, Juan Fernandez); Lasiococca (5; L. brevipes, L. chanii, L. comberi, L. locii, L. symphyllifolia; eastern Himalayas, Hainan, Vietnam, the Malay Peninsula), Spathiostemon (2; S. javensis, S. moniliformis; Peninsular Thailand, Malesia to New Guinea), Homonoia (3; H. intermedia, H. retusa, H. riparia; Malesia New Guinea); Lobanilia (8; Madagascar); Macaranga (305–310; tropical regions in the Old World); Mareya (4; M. acuminata, M. brevipes, M. congolensis, M. micrantha; tropical West and Central Africa); Mercurialis (12; Europe, the Mediterranean, temperate Asia to northern Thailand), Seidelia (2; S. firmula, S. triandra; Northern and Western Cape, Free State), Leidesia (1; L. procumbens; southern Africa); Ricinus (1; R. communis; eastern and northeastern Africa, the Arabian Peninsula, southwestern Asia), Adriana (2; A. quadripartita, A. urticoides; Australia); Avellanita (1; A. bustillosii; central Chile), Mallotus (120–125; tropical Asia to tropical Australia and islands in the Pacific, two species, M. oppositifolius and M. subulatus, in tropical Africa and Madagascar); Blumeodendron (5; B. bullatum, B. calophyllum, B. concolor, B. kurzii, B. tokbrai; Southeast Asia, the Andaman Islands, Malesia to New Guinea), Podadenia (1; P. sapida; Sri Lanka), Ptychopyxis (13; Peninsular Thailand, Malesia to New Guinea), Botryophora (1; B. geniculata; Southeast Asia, West Malesia); Afrotrewia (1; A. kamerunica; Cameroon, Gabon). – Adelieae G. L. Webster in Taxon 24: 597. 19 Dec 1975. Adelia (10; Mexico, Central America, the West Indies, tropical South America), Enriquebeltrania (2; E. crenatifolia, E. disjuncta; Mexico), Garciadelia (4; G. abbottii, G. castilloae, G. leprosa, G. mejiae; Hispaniola), Lasiocroton (c 25?; Cuba), Leucocroton (26–27; Cuba). – Agrostistachydeae G. L. Webster in Taxon 24: 596. 19 Dec 1975. Agrostistachys (6; A. borneensis, A. gaudichaudii, A. hookeri, A. indica, A. sessilifolia, A. staminodiata; southern India, Sri Lanka, Thailand, Malesia to New Guinea), Chondrostylis (2; C. bancana, C. kunstleri; Peninsular Thailand, West Malesia), Cyttaranthus (1; C. congolensis; Central Africa), Pseudagrostistachys (2; P. africana, P. ugandensis; tropical Africa). – Alchorneeae Hutch. in Amer. J. Bot. 56: 752. Aug 1969. Alchornea (50–55; tropical regions on both hemispheres), Aparisthmium (1; A. cordatum; southern Central America, tropical South America), Bocquillonia (14; New Caledonia), Orfilea (4; O. ankafinensis, O. coriacea, O. multispicata, O. neraudiana; Madagascar, Mauritius); Aubletiana (2; A. leptostachys, A. macrostachys; Cameroon, Gabon), Conceveiba (14; Central America, tropical South America), Mareyopsis (2; M. longifolia, M. oligogyna; Central Africa). – Ampereae Müll.-Arg. in Bot. Zeitung (Berlin) 22: 324. 14 Oct 1864. Monotaxis (11; Australia), Amperea (8; southwestern Western Australia, southeastern Australia, Tasmania). – Bernardieae G. L. Webster in Taxon 24: 596. 19 Dec 1975. Bernardia (c 75; southern United States, Mexico, Central America, tropical South America, with their highest diversity in Brazil), Discocleidion (1; D. rufescens; central China, the Ryukyu Islands), Adenophaedra (3; A. cearensis, A. grandifolia, A. megalophylla; tropical South America). – Caryodendreae G. L. Webster in Taxon 24: 596. 19 Dec 1975. Caryodendron (4; C. amazonicum, C. angustifolium, C. janeirense, C. orinocense; Costa Rica to Amazonian Brazil), Discoglypremna (1; D. caloneura; tropical Africa), Alchorneopsis (2; A. floribunda; Central America, tropical South America, one species, A. portoricensis, on Puerto Rico). – Chrozophoreae Pax et K. Hoffm. in H. G. A. Engler, Pflanzenr. 68(Addit. VI): 2. 6 Jun 1919. Chrozophora (11; the Mediterranean, tropical East Africa, southwestern and southern Asia to Thailand); Argythamnia (23; southern Mexico to Honduras, the West Indies), Caperonia (c 35; tropical Africa, Madagascar, tropical America), Chiropetalum (c 25; Texas, Mexico, tropical South America to northern Chile), Ditaxis (c 50; Florida, southwestern United States, Mexico, Central America, the West Indies, tropical South America), Philyra (1; P. brasiliensis; southern Brazil, Paraguay, northern Argentina); Doryxylon (1; D. spinosum; Luzon, the Lesser Sunda Islands), Melanolepis (2; M. multiglandulosa, M. vitifolia; Southeast Asia to islands in the Pacific, Taiwan), Sumbaviopsis (1; S. albicans; Assam, Southeast Asia, West Malesia), Thyrsanthera (1; T. suborbicularis; Southeast Asia); Speranskia (3; S. cantonensis, S. tuberculata, S. yunnanensis; southern China, northern Burma). – Epiprineae Hurus. in J. Fac. Sci. Univ. Tokyo, ser. 3, Bot. 6: 309. 1954. Epiprinus (6; E. balansae, E. lanceifolius, E. malayanus, E. mallotiformis, E. poilanei, E. siletianus; India, Burma, southern China, Southeast Asia, Sumatra), Cleidiocarpon (2; C. cavaleriei, C. laurinum; Burma, southern China, western Thailand, northern Vietnam), Koilodepas (12; southern India to Hainan and New Guinea), Cladogynos (1; C. orientalis; Southeast Asia, Malesia), Tsaiodendron (1; T. dioicum; Yunnan), Cephalocroton (4; C. cordofanus, C. incanus, C. mollis, C. polygynus; tropical Africa, Madagascar, Socotra, Sri Lanka); Cephalomappa (5; C. beccariana, C. lepidotula, C. malloticarpa, C. paludicola, C. penangensis; southern China, West Malesia), Rockinghamia (2; R. angustifolia, R. brevipes; northeastern Queensland). – Erismantheae G. L. Webster in Taxon 24: 595. 19 Dec 1975. Erismanthus (2; E. obliquus, E. sinensis; Southeast Asia, Hainan, West Malesia); Moultonianthus (1; M. leembruggianus; Sumatra, Borneo), Syndyophyllum (2; S. occidentale: northern Sumatra, Borneo; S. excelsum: northern Papua New Guinea). – Plukenetieae Hutch. in Amer. J. Bot. 56: 753. Aug 1969. Dalechampia (120–125; tropical regions on both hemispheres, with their highest diversity in tropical South America); Haematostemon (2; H. coriaceus, H. guianensis; Guyana, Amazonian Venezuela), Plukenetia (c 20; tropical Africa, Madagascar, southern Mexico, Central America, tropical South America, one species, P. corniculata, in tropical Asia), Romanoa (1; R. tamnoides; eastern and southern Brazil, Paraguay, Bolivia); Ctenomeria (2; C. capensis, C. cordata; South Africa), Megistostigma (5; M. burmanicum, M. cordatum, M. glabratum, M. peltatum, M. yunnanense; Yunnan, Southeast Asia, West Malesia), Cnesmone (11; Assam, Southeast Asia, West Malesia), ‘Tragia’ (150–155; tropical, subtropical and warm temperate regions on both hemispheres; polyphyletic), Bia (5; B. alienata, B. cordata, B. fallax, B. fendleri, B. lessertiana; Costa Rica to tropical South America), Platygyna (7; P. dentata, P. hexandra, P. leonis, P. obovata, P. parvifolia, P. triandra, P. volubilis; Cuba), Acidoton (6; A. haitiensis, A. lanceolatus, A. microphyllus, A. nicaraguensis, A. urens, A. variifolius; Hispaniola and Jamaica), Zuckertia (2; Z. cordata, Z. manuelii; Mexico, Central America), ‘Tragia’ pro parte, Pachystylidium (1; P. hirsutum; India, Southeast Asia to Central Malesia), Sphaerostylis (2; S. perrieri, S. tulasneana; Madagascar), Gitara (1; G. nicaraguensis; Central America to Venezuela), ‘Tragia’ pro parte; unplaced Plukenetieae: Angostylis (2; A. longifolia, A. tabulamontana; Amazonian Brazil), Astrococcus (1; A. cornutus; Amazonian Venezuela, Amazonian Brazil). – Pycnocomeae Reveal in Phytoneuron 2012-37: 218. 23 Apr 2012. Amyrea (11; Madagascar), Necepsia (3; N. afzelii, N. castaneifolia, N. zairensis; tropical Africa, Madagascar), Paranecepsia (1; P. alchorneifolia; tropical East Africa); Pycnocoma(18; tropical Africa), Droceloncia (1; D. rigidifolia; Madagascar, the Comoros), Argomuellera (17; tropical Africa, Madagascar). – Sphyranthereae Radcl.-Sm., Gen. Euphorbiacearum: 135. 2001. Sphyranthera (2; S. airyshawii, S. lutescens; the Andaman Islands, the Nicobar Islands). – Pantropical, few species in subtropical and warm-temperate regions. Pseudanthia present in Dalechampia. Stigma in Acalypha strongly branched. Outer integument three to six (to 16) cell layers thick. Inner integument three to 24 cell layers thick. Testa sometimes with vascular bundles.

[Crotonoideae+Euphorbioideae]

Laticifers and latex present. Large variation among phorbol esters; cocarcinogens present.

Crotonoideae (Kunth) Beilschm. in Flora 16(Beibl. 7): 61, 106. 14 Jun 1833 [’Crotoneae’]

60/2.010–>2.020. Aleuritideae Hurus. in J. Fac. Sci. Univ. Tokyo, ser. 3, Bot., 6: 309. 15 Aug 1954. Aleurites (2; A. moluccanus, A. rockinghamensis; tropical Asia to islands in western Pacific), Vernicia (3; V. cordata, V. fordii, V. montana; Burma, Southeast Asia, Malesia and southern China to Japan); Benoistia (3; B. orientalis, B. perrieri, B. sambiranensis; Madagascar); Cyrtogonone (1; C. argentea; tropical West Africa), Crotonogyne (16; tropical Africa), Manniophyton (1; M. fulvum; western and central tropical Africa to Angola); Garcia (2; G. nutans, G. parviflora; Mexico to Colombia); Grossera (8; tropical Africa, Madagascar), Cavacoa (3; C. aurea, C. baldwinii, C. quintasii; tropical Africa), Sandwithia (2; S. guyanensis, S. heterocalyx; northeastern South America), Tannodia (9; tropical Africa, Madagascar), Tapoides (1; T. villamilii; Borneo); Neoboutonia (3; N. macrocalyx, N. mannii, N. melleri; tropical Africa); Deutzianthus (2; D. tonkinensis; northern Vietnam; D. thyrsiflorus: Sumatra), Oligoceras (1; O. eberhardtii; Vietnam), Paracroton (4; P. integrifolius, P. pendulus, P. sterrhopodus, P. zeylanicus; southern India, Sri Lanka, Malesia to New Guinea); Mildbraedia (5; M. balboana, M. carpinifolia, M. fallax, M. occidentalis, M. paniculata; western and central tropical Africa to Mozambique). – Codiaeeae Hutch. in Amer. J. Bot. 56: 747. Aug 1969 [‘Codiaeae’]. Alphandia (3; A. furfuracea, A. resinosa, A. verniciflua; New Guinea, New Caledonia, Vanuatu), Anomalocalyx (1; A. uleanus; Amazonian Brazil from near Manaos to Amapá), Baliospermum (5; B. angustifolium, B. bilobatum, B. calycinum, B. solanifolium, B. yui; the Himalayas, Tibet, Yunnan, Indochina, the Malay Peninsula, Sumatra, Java, Sumbawa), Baloghia (15; eastern Queensland, eastern New South Wales, Norfolk Island, New Caledonia, with the largest diversity in New Caledonia), Blachia (11; India, the Andaman Islands, southern China, Southeast Asia, the Philippines), Codiaeum (16–17; Malesia to New Guinea, tropical Australia and New Caledonia), Dimorphocalyx (17; India, Sri Lanka to the Philippines, New Guinea and tropical Australia), Dodecastigma (3; D. amazonicum, D. integrifolium, D. uleanum; the Guianas, Amazonian Brazil), Fontainea (9; New Guinea, Queensland, New Caledonia, Vanuatu), Hylandia (1; H. dockrillii; northeastern Queensland), Ostodes (2; O. kuangii, O. paniculata; Assam, eastern Himalayas, Southeast Asia to Java and Borneo), Pantadenia (3; P. chauvetiae and P. gervaisii: Madagascar; P. adenanthera: Thailand, Indochina), Strophioblachia (1; S. fimbricalyx; Yunnan, Hainan, Thailand, Cambodia, Vietnam, the Philippines, Sulawesi); Trigonostemon (c 85; India and China to the Philippines, New Guinea, Queensland and Fiji); Pausandra (8; Central America, tropical South America). – Crotoneae Dumort., Anal. Fam. Plant.: 45. 1829 [‘Crotonieae’]. Acidocroton (13; southern Mexico, Central America to Colombia, Cuba, Hispaniola, Jamaica), Brasiliocroton (1; B. mamoninha; northeastern Brazil), Croton (>1.200; tropical and subtropical regions), Moacroton (>28; tropical and subtropical North, Central and South America, the West Indies), Sagotia (2; S. brachysepala, S. racemosa; Central America, northern South America), Sandwithia (2; S. guyanensis, S. heterocalyx; Amazonian South America). – Elateriospermeae G. L. Webster in Taxon 24: 599. 19 Dec 1975. Elateriospermum (1; E. tapos; Peninsular Thailand, West Malesia), Glycydendron (2; G. amazonicum, G. espiritosantense; northern South America). – Heveeae (Müll. Arg.) G. L. Webster in Kubitzki (ed) 2014 Fam. Gen. Vasc. Plants, p. 164. Hevea (9; the Amazon basin). – Jatropheae Baill., Hist. Plant. 5: 156, 179. Jan–Apr 1874. Jatropha (>190; tropical and subtropical regions on both hemispheres, North America), Joannesia (2; J. heveoides, J. princeps; Venezuela, Amazonian and coastal Brazil), Vaupesia (1; V. cataractarum; Colombia, western Brazil). – Manihoteae Pax in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. III, 5: 14. Mai 1890. Cnidoscolus (c 95; southern United States, Mexico, Central America, the West Indies, tropical South America), Manihot (105–110; southwestern United States, Mexico, Central America, the West Indies, tropical South America). – Micrandreae G. L. Webster in Taxon 24: 598. 19 Dec 1975. Hevea (c 10; the Amazon basin); Micrandra (11–12; tropical South America), Micrandropsis (1; M. scleroxylon; Amazonas, Amazonian Colombia). – Ricinocarpeae Müll.-Arg. in Bot. Zeitung (Berlin) 22: 324. 14 Oct 1864. Bertya (28; southern and eastern Australia), Borneodendron (1; B. aenigmaticum; ultrabasic soils on northern Borneo), Cocconerion (2; C. balansae, C. minus; ultrabasic soils in New Caledonia), Myricanthe (1; M. discolor; ultrabasic soils in northwestern New Caledonia); Beyeria (c 25; southern and eastern Australia, Tasmania), Ricinocarpos (28; western and northern Australia, New Caledonia), Shonia (4; S. bickertonensis, S. carinata, S. territorialis, S. tristigma; Northern Territory, Queensland). – Ricinodendreae Hutch. in Amer. J. Bot. 56: 749. Aug 1969. Annesijoa (1; A. novoguineensis; New Guinea), Givotia (4; G. gosai, G. madagascariensis, G. moluccana, G. stipularis; tropical East Africa, Madagascar, India, Sri Lanka), Leeuwenbergia (2; L. africana, L. letestui; Gabon to Cameroon and Congo), Ricinodendron (2; R. heudelotii, R. lobatus; tropical Africa to Angola and Mozambique); incertae sedis: Radcliffea (1; R. smithii; western Madagascar). – Pantropical, few species in East Asia and North America. Laticifers articulated or inarticulated. Hairs often stellate or lepidote. Lamina sometimes with abaxial paired glands near petiole junction. Petals present or absent. Pollen grains colpate, porate or inaperturate. Seeds often pachychalazal. Aril or carunculus often present. Exotesta sometimes palisade. Endotestal cells sometimes palisade, thin-walled, slightly lignified. Tegmen usually vascularized. Cyanogenes via valine/isoleucine pathway. Deletion of more than 100 bp in plastid trnL/F spacer in some species.

Euphorbioideae (Kunth) Beilschm. in Flora 16(Beibl. 7): 61, 105. 14 Jun 1833 [‘Euphorbieae’]

38/2.410–2.420. Euphorbieae Dumort., Anal. Fam. Plant.: 45. 1829. Anthostema (3; A. aubryanum, A. madagascariense, A. senegalense; tropical West Africa, Madagascar), Dichostemma (2; D. glaucescens: Central Africa; D. zenkeri: Cameroon); Neoguillauminia (1; N. cleopatra; New Caledonia), Calycopeplus (5; C. casuarinoides, C. collinus, C. marginatus, C. oligandrus, C. paucifolius; northern Australia, southwestern Western Australia); Euphorbia (c 2.050; almost cosmopolitan). – Hippomaneeae Bartl., Ord. Nat. Plant.: 372. Sep 1830 [‘Hippomanea’]. Homalanthus (23; tropical Asia to New Guinea, northern and eastern Australia, New Caledonia, New Zealand, Polynesia); Colliguaja (5; C. brasiliensis, C. dombeyana, C. integerrima, C. odorifera, C. salicifolia; southern Brazil, central Chile and adjacent Paraguay and Uruguay), Grimmeodendron (2; G. eglandulosum, G. jamaicense; the West Indies), Bonania (7; B. cubana, B. domingensis, B. elliptica, B. emarginata, B. erythrosperma, B. linearifolia, B. myricifolia; the West Indies), Adenopeltis (1; A. serrata; Peru, Chile), Stillingia (c 30; Madagascar, the Mascarene Islands, East Malesia, Fiji, southern United States, Mexico, Central America, the West Indies, tropical South America), Gradyana (1; G. franciscana; northeastern Brazil), Spegazziniophytum (1; S. patagonicum; southern Argentina), Sapium (23; Mexico, Central America, the West Indies, tropical South America), Hippomane (3; H. horrida, H. spinosa: Hispaniola; H. mancinella: Florida to Venezuela, the Galápagos Islands), Senefelderopsis (2; S. chiribiquetensis, S. croizatii; the Guayana Highlands and adjacent areas), Incadendron (1; I. esseri; Ecuador, Peru), Pleradenophora (1; P. longicuspis; Mexico, Guatemala, Belize), Balakata (2; B. baccata, B. luzonica; southern China, tropical Asia, New Guinea, tropical Australia), Falconeria (1; F. insignis; the Himalayas, Southeast Asia, West Malesia), Sclerocroton (6; S. carterianus, S. cornutus, S. integerrimus, S. melanostictus, S. oblongifolius, S. schmitzii; tropical Africa, Madagascar), Triadica (3; T. cochinchinensis, T. rotundifolia, T. sebifera; East and tropical Asia), Mabea (c 40; Mexico, Central America, tropical South America), Gymnanthes (c 25; tropical and southern Africa, tropical Asia to Malesia and islands in the Pacific, Florida, Mexico, Central America, the West Indies, tropical South America, one species, G. japonica, in central China, the Korean Peninsula and Japan), Actinostemon (c 20; the West Indies, tropical South America), Senefeldera (6; S. inclinata, S. macrophylla, S. multiflora, S. testiculata, S. triandra, S. verticillata; tropical South America), Dalembertia (4; D. hahniana, D. platanoides, D. populifolia, D. triangularis; Mexico, Guatemala), Maprounea (6; M. africana, M. amazonica, M. brasiliensis, M. guianensis, M. membranacea, M. obtusata; tropical Africa, tropical South America), Excoecaria (c 40; Africa to Australia and Melanesia), Sebastiania (70–75; southwestern United States, Mexico, Central America, the West Indies, tropical South America), Algernonia (12; Peru, eastern Brazil), Hura (2; H. polyandra: Mexico, Central America; H. crepitans: Nicaragua to the West Indies, Peru and Brazil), Ophthalmoblapton (4; O. crassipes, O. macrophyllum, O. parviflorum, O. pedunculare; eastern Brazil); Pachystroma (1; P. longifolium; southeastern Brazil, Bolivia, Peru). – Stomatocalyceae G. L. Webster in Taxon 24: 600. 19 Dec 1975. Plagiostyles (2; P. africana, P. pinnatus; southern Nigeria, Gabon, Congo), Pimelodendron (4; P. amboinicum, P. griffithianum, P. macrocarpum, P. zoanthogyne; Malesia to New Guinea, northeastern Queensland); Hamilcoa (1; H. zenkeri; Nigeria, Cameroon), Nealchornea (2; N. stipitata, N. yapurensis; Colombia, Brazil, eastern Peru). – Subcosmopolitan. Laticifers inarticulated. White caustic latex present. Starch grains often complex in latex (e.g. in Euphorbia). Disc usually absent. Stamens not covered by tepals. Outer integument three to six or eight to 22 cell layers thick. Inner integument three to five (to 22) cell layers thick. Carunculus often present.

Cladogram of Euphorbiaceae based on DNA sequence data (Tokuoka 2007).

Vegetative anatomy Phellogen? Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits absent. Imperforate tracheary xylem elements tracheids? or libriform fibres with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal aliform, winged-aliform, or confluent. Sieve tube plastids S type? Nodes? Cortex and medulla with sclereids.

Leaves Alternate (spiral), simple, entire, coriaceous, with revolute ptyxis. Stipules small; leaf sheath absent. Petiole vascular bundle transection annular; bundles not always immediately fused with stele. Lamina densely tomentose on abaxial side. Venation pinnate. Stomata paracytic? Cuticular wax crystalloids? Hypodermis with mucilaginous idioblasts. Mesophyll with sclerenchymatous idioblasts. Leaf margin entire, with abaxial glands at base.

Flowers Obliquely zygomorphic. Half epigyny. Sepals five, non-uniform, with imbricate quincuncial aestivation, connate at base into short hypanthium-like structure (“floral cup”). Petals three (abaxial-lateral and abaxial petals absent), with contorted aestivation, free. Nectariferous disc annular, intrastaminal, inserted at adaxial side of hypanthium-like structure. Calcium oxalate druses present in floral parts.

Androecium Stamens usually four fertile (sometimes five to seven): two longer outer antesepalous and two shorter inner alternisepalous, separated by one long acute abaxial-lateral antesepalous staminodium on one side and (one to) three to five short dentate adaxial staminodia on opposite side. Filaments connate at base into two groups, adnate to petals (epipetalous). Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often thickened dorsally? Tapetum secretory? Staminodia two to six.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine?, with ? infratectum, sculpturing?

Gynoecium Pistil composed of three connate carpels; median carpel adaxial; carpel semi-synascidiate. Ovary semi-inferior, trilocular, with central columella; locules filled by unicellular hairs. Style single, simple. Stigma capitate, papillate, probably Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules two per carpel, anatropous, apotropous, upper ovule with micropyle directed downwards, lower ovule with micropyle directed upwards, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument three or four cell layers thick. Inner integument six or seven cell layers thick. Obturator probably absent. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule with thin exocarp, persistent calyx and persistent central columella.

Seeds Aril absent. Testa winged. Tegmen? Perisperm not developed. Endosperm thin and sparse or absent. Embryo straight?, chlorophyll? Cotyledons two? Germination phanerocotylar?

Vegetative anatomy Phellogen ab initio superficial. Vascular cylinder and medulla quadrangular or quinquangular. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Wood fluorescent. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Heartwood with gum-like substances. Prismatic calciumoxalate crystals and druses present.

Trichomes Hairs unicellular or multicellular, uniseriate, simple, with thickened cell walls and pitted base.

Leaves Alternate (distichous), simple, entire, coriaceous, with ? ptyxis. Stipules narrowly elongate, inflexed, caducous; leaf sheath absent. Petiole vascular bundle transection annular; petiole with inverted medullary bundle. Venation pinnate to palmate, acrodromous (actinodromous); secondary veins ascending; tertiary veins scalariform. Stomata usually laterocytic (rarely anisocytic). Cuticular waxes absent. Abaxial domatia present in vein axils. Sclereids simple or branched. Epidermal cells mucilaginous. Leaf margin serrate or entire.

Flowers Actinomorphic. Pedicel articulated? Hypogyny. Sepals five, with imbricate aestivation, connate (free?). Petals five, with induplicate-valvate aestivation, long and subulate, apically inflexed in bud (sometimes geniculate or sigmoid at anthesis), free. Nectariferous disc annular, thin, intrastaminal, sinuate at margin.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments very short, free from each other and from tepals, inserted at margin of nectariferous disc. Anthers basifixed, adnate (locules short, somewhat separate), non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective somewhat extended, with long apical hairs. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate or tricolpate, shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate? infratectum, reticulate. Endexinal folds present.

Gynoecium Pistil composed of five connate antepetalous carpels. Ovary superior, quinquelocular. Stylodia five, separate, short, with adaxial furrows, inserted at outer carpellary margins. Stigmas five, subulate, type? Pistillodium absent.

Ovules Placentation basal to axile. Ovules few per carpel, anatropous, ascending, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument approx. three cell layers thick. Inner integument approx. three cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Testa reticulate, with mesotesta consisting of sclereids. Endotesta? Exotegmen fibrous, little developed, ridged, sclereidal; exotegmic cells with U-shaped wall thickenings. Endotegmen? Perisperm not developed. Endosperm copious, fleshy. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Distribution Southern Mexico, Central America, tropical South America; one species of Sacoglottis in tropical West Africa.

Fossils Uncertain. Fossil endocarps sometimes assigned to Sacoglottis and Vantanea have been found in Miocene and Pliocene layers from Costa Rica in the north to Bolivia in the south (Herrera & al. 2010).

Habit Bisexual, evergreen trees or shrubs. Young branches angular in cross-section. Often with aromatic juice.

Vegetative anatomy Phellogen ab initio subepidermal. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal aliform, lozenge-aliform, winged-aliform, vasicentric, or unilateral. Tyloses sometimes abundant. Sieve tube plastids Pcs type, with protein crystals and starch. Nodes 3:3, trilacunar with three leaf traces. Mucilage cells numerous. Secretory cavities absent. Heartwood sometimes with gum-like substances. Silica bodies present in some species. Prismatic calciumoxalate crystals abundant.

Leaves Alternate (spiral or distichous), simple, entire, coriaceous, usually with involute ptyxis. Stipules small and caducous, or absent; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata paracytic or anomocytic. Cuticular wax crystalloids? Secretory cavities absent. Mesophyll with or without sclerenchymatous idioblasts (with sclereids reaching from one epidermis to opposite side of lamina) and with calciumoxalate as druses or single prismatic crystals. Leaf margin usually serrate (sometimes crenate or entire). Extrafloral nectaries rarely present on adaxial side of lamina (e.g. in Vantanea).

Flowers Actinomorphic or somewhat zygomorphic. Hypogyny. Sepals (four or) five, two outer ones often smaller, with imbricate aestivation, persistent, connate in lower part into tube. Petals (four or) five, with usually quincuncial to cochlear (sometimes imbricate or contorted) aestivation, thick, persistent or caducous, free. Nectariferous disc intrastaminal, with stomata, usually cupular to tubular or dentate to lobate (sometimes as ten to 20 free scales), often adnate to ovary base or filament bases.

Androecium Stamens usually ten to c. 30, in one to five whorls (sometimes in five antesepalous staminal fascicles each with three stamens, and five antepetalous stamens; in Vantanea c. 40 to more than 100 stamens in fascicles); filament bundles alternipetalous. Filaments connate into tube at least in lower part, free from tepals. Anthers dorsifixed or subbasifixed or inserted at connective bases, versatile, (disporangiate or) tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits), with thecae separated, superposed; connective wide, apically prolongate. Tapetum secretory. Staminodia present in some species.

Pollen grains: Microsporogenesis simultaneous. Pollen grains usually 3(–4)-colporate (rarely 3–4-porate), shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, microreticulate, perforate or punctate.

Gynoecium Pistil composed of (four or) five (to seven) connate usually antesepalous carpels. Ovary superior, (quadrilocular or) quinquelocular (to septalocular; sometimes unilocular at apex). Style single, simple. Stigma entire to slightly lobate, type? Pistillodium absent.

Ovules Placentation apical to axile. Ovules one (or two) per carpel, anatropous, pendulous, epitropous (micropyle directed upwards-outwards), bitegmic, crassinucellar. Micropyle usually bistomal or exostomal (rarely endostomal). Outer integument usually two cell layers thick, tanniniferous. Inner integument three cell layers thick, tanniniferous. Endothelium absent. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A one- or two-seeded drupe, with uneven surface and operculate multilocular pyrene, sometimes with numerous cavities filled with resinous? material (for floating on water surface).

Seeds Aril absent. Exotestal cell walls thick and lignified. Endotesta? Tegmen multiplicative (approx. five cell layers thick). Exotegmen fibrous; cross layer present beneath exotegmen. Endotegmen? Perisperm not or sparsely developed. Endosperm copious, oily. Embryo straight or slightly curved, well differentiated, chlorophyll? Cotyledons two. Germination?

Phytochemistry Very insufficiently known. Ellagic acid present. Bergenin and gallate present in bark in Sacoglottis. Alkaloids not found.

Ascyraceae Plenck, Elem. Termin. Bot.: 162. 1796 [’Asciroideae’]; Hypericales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 218. Jan-Apr 1820 [‘Hypericinae’]

Distribution Temperate regions on both hemispheres, tropical mountains, tropical regions in Africa, Madagascar and America.

Habit Bisexual, evergreen or deciduous trees, shrubs or herbs. Species in dry areas sometimes with lignotuber.

Vegetative anatomy Phellogen ab initio superficial or pericyclic. Polyderm often present. Endodermis sometimes prominent. Vessel elements usually with simple (sometimes scalariform? or opposite) perforation plates; lateral pits alternate, bordered pits present. Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits (also vasicentric tracheids), septate (Hypericum) or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma paratracheal scanty, vasicentric, reticulate, or banded, or absent. Tyloses sometimes abundant. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Schizogenous glands, canals and cavities with resin, balsam or yellow to red secretions with hypericin and closely allied compounds. Colleters often present. Silica bodies or prismatic calciumoxalate crystals present in some species.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, often dendritic or stellate.

Leaves Usually opposite (rarely verticillate or alternate), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Colleters often present. Petiole vascular bundle transection arcuate, annular or complex. Venation pinnate, usually eucamptodromous or brochidodromous. Stomata usually paracytic (sometimes anomocytic, anisocytic or cyclocytic). Cuticular wax crystalloids as parallel grouped (usually non-entire) platelets (Hypericum type). Lamina with glandular dots or glandular lines (pellucid-punctate dots, resin/latex cavities). Idioblasts or schizogenous glands and canals and cavities present. Leaf margin usually entire (rarely serrate).

Inflorescence Usually terminal (rarely axillary), cymose (often thyrsoid, scorpioid?), or flowers solitary.

Flowers Actinomorphic. Hypogyny. Sepals (two to) four or five, with decussate or quincuncial aestivation, free. Petals (three or) four or five, often with contorted aestivation, free. Nectary absent? Disc absent.

Androecium Stamens nine to more than 650, centrifugally developing. Filaments thin, free or often connate at base into three to five antepetalous fascicles, free from tepals. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, usually introrse (sometimes extrorse), longicidal (dehiscing by longitudinal slits); connective often with apical glands. Tapetum secretory. Staminodia three or five (nectaries?), alternipetalous, or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tri(col)porate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or microreticulate, sometimes psilate.

Gynoecium Pistil composed of three to five connate antesepalous carpels. Ovary superior, trilocular to quinquelocular. Stylodia usually three to five, free or connate at base. Stigmas expanded, punctate to widened, papillate or non-papillate, usually Dry type. Pistillodium absent.

Ovules Placentation usually axile (sometimes parietal). Ovules one to numerous per carpel, anatropous, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument up to seven cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus? Antipodal cells often proliferating (in Hypericum; up to seven cells). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad.

Fruit Usually a loculicidal and/or septicidal capsule or a berry (rarely a drupe).

Seeds Aril absent. Seeds sometimes winged. Operculum often? present. Seed coat exotegmic. Testa sometimes glandular. Exotesta often with tanniniferous epidermal cells. Endotesta? Exotegmen palisade, with anticlinal cell walls sinuate, low and lignified (sometimes absent). Endotegmen? Perisperm not developed. Endosperm sparse or absent. Embryo usually straight (fusiform, sometimes curved), rudimentary or well differentiated, with or without chlorophyll. Cotyledons two, often large. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavones, biflavonoids, xanthones (e.g. mangiferin), benzophenones, and polyacetate-derived and other anthraquinones (vismiones) present. In Hypericum also emodin derivatives, prenylated phloroglucinol derivatives (hyperforin), and naphthodianthrones (pseudohypericin and hypericin, possibly synthesized by endophytic relative to Chaetomium). In Vismieae also anthrones, biemodyles and closely allied compounds. Ellagic acid not found.

Distribution Tropical West and Central Africa, Madagascar, Southeast Asia, Malesia.

Fossils The fossil wood Irvingiaceoxylon dechampsii was reported from the Upper Pliocene/Lower Pleistocene of Ethiopia (Gros 1983).

Vegetatively anatomy Phellogen ab initio deeply seated. Vessel elements with simple perforation plates; lateral pits alternate, simple pits? Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays uniseriate, homocellular. Axial parenchyma apotracheal diffuse?, or paratracheal aliform, winged-aliform, confluent, reticulate, or banded. Tyloses abundant. Sieve tube plastids S type. Nodes trilacunar? Secretory cavities with mucilage; parenchyma without secretory cavities (secretory ducts absent from stem in Allantospermum). Prismatic calciumoxalate crystals often frequent.

Leaves Alternate (distichous), simple, entire, coriaceous, with revolute ptyxis. Stipules large, intrapetiolar, enclosing terminal bud, early caducous; leaf sheath absent. Petiole vascular bundle transection annular. Venation pinnate; secondary veins subparallel; tertiary veins parallel. Stomata paracytic or anomocytic. Cuticular wax crystalloids? Epidermis with mucilaginous idioblasts. Mesophyll with mucilaginous idioblasts and canals, and with sclerenchymatous idioblasts containing calciumoxalate druses or solitary prismatic crystals. Cristarque cells abundant. Sclereids present in Irvingia. Leaf margin entire.

Flowers Actinomorphic, small. Pedicel articulated at base. Hypogyny. Sepals five, with imbricate aestivation, recurved, persistent, free. Petals five, with cochlear or quincuncial aestivation, free. Nectariferous disc intrastaminal, massive, lobate.

Androecium Stamens (nine or) ten, not more than twice as many as petals, diplostemonous. Filaments incurved-folded in bud, inserted below nectariferous disc, free from each other and from tepals. Anthers basifixed to slightly dorsifixed, versatile, tetrasporangiate, introrse (antesepalous stamens) or almost latrorse (antepetalous stamens), longicidal (dehiscing by longitudinal slits); connective not protruding. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular at dispersal. Tectum continuous. Exine?, with ? infratectum, verrucate or striate-rugulate

Gynoecium Pistil composed of two (Allantospermum, Desbordesia), four or five connate carpels; carpels median (when two) or antesepalous. Ovary superior, bilocular, quadrilocular or quinquelocular, synascidiate. Style single, simple, short. Stigma capitate, unicellular-papillate, type? Pistillodium absent.

Ovules Placentation apical to axile. Ovule one per carpel, anatropous to hemianatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument three cell layers thick. Inner integument three or four cell layers thick, non-multiplicative. Integumentary tapetum absent. Obturator placental. Endothelium absent. Megasporangium with cytoplasm-rich peripheral cell layers filled with starch grains. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A drupe with one, four or five single-seeded (or five-seeded) pyrenes (often with radial fibres) or a bilocular samara (Desbordesia) with persistent calyx.

Seeds Aril absent. Seed coat testal. Testa thick, strongly sclerotic, multiplicative, highly vascularized. Tegmen non-multiplicative. Exotegmen fibrous. Perisperm not developed. Endosperm sparse to copious. Embryo?, chlorophyll? Cotyledons two, large, cordate. Germination phanerocotylar, epigeal.

Phytochemistry Very insufficiently known. Ellagic and gallic acids present. Seed lipids with myristic acid and lauric acid.

Miquel, Fl. Ned. Ind. 1(2): viii, 494. 30 Sep 1858 [‘Ixionantheae’], nom. cons.

Distribution Tropical Africa, northeastern India, eastern Himalayas, southern China, Southeast Asia, Malesia to New Guinea, northeastern South America.

Vegetative anatomy Phellogen? Primary medullary strands narrow and wide, alternating. Vessel elements with simple perforation plates; lateral pits alternate, scalariform or opposite, simple or bordered pits. Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse-in-aggregates, or paratracheal scanty, reticulate, or banded. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Mucilage cells and mucilage cavities absent. Tracheoidal idioblasts often present. Most species with secretory ducts in stem? Silica bodies sometimes abundant. Prismatic calciumoxalate crystals abundant.

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, with involute ptyxis. Stipules small lateral (sometimes cauline); leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids as platelets arranged in various ways. Mesophyll in Ochthocosmus with sclerenchymatous idioblasts (some sclereids with spiral thickenings). Leaf margin usually serrate (rarely entire). Extrafloral nectaries sometimes present on lamina.

Flowers Actinomorphic, small. Half epigyny. Sepals (four or) five, with imbricate or contorted aestivation, usually connate at base. Petals (four or) five, with imbricate or contorted aestivation, often marcescent, free. Nectariferous disc intrastaminal, usually annular or cupular, free (in Ochthocosmus as interstaminal glands).

Androecium Stamens (four or) five, antesepalous, or up to 20 (antepetalous stamens paired in Ixonanthes, arising from common strand). Filaments widened at base, sigmoid-folded in bud, free from each other and from tepals, free from or adnate at base to nectariferous disc. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, spinulate.

Gynoecium Pistil composed of usually five (rarely two or four) connate carpels; carpels sometimes divided. Ovary semi-inferior, sometimes unilocular at apex; locules sometimes divided into locelli by incomplete secondary septa. Style single, simple, filiform, folded in bud. Stigma single, capitate to discoid, type? Pistillodium absent.

Ovules Placentation apical to axile. Ovules usually two per carpel, anatropous, pendulous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument approx. four cell layers thick. Obturator placental. Hypostase present. Endothelium present. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit Usually a septicidal (rarely also loculicidal, through secondary septa) capsule, also dehiscing adaxially, with persistent sepals and petals and, sometimes, persistent central columella.

Seeds Aril present between hilum and micropyle, or testa winged at base (Ixonanthes, Ochthocosmus). Exotegmen fibrous. Endotegmic cells with sinuous anticlinal walls. Perisperm not developed. Endosperm usually sparse or absent. Embryo straight, chlorophyll? Cotyledons two, large. Germination?

Phytochemistry Very insufficiently known. Ellagic acid and proanthocyanidins present. Alkaloids not found.

von Martius, Nov. Gen. Sp. Plant. 1: 154, 158. Jan-Mar 1826 [’Lacistemeae’], nom. cons.

Lacistematales Mart. in C. F. P. von Martius, Consp. Regn. Veg.: 49. Sep-Oct 1835 [’Lacistemeae’]

Distribution Southern Mexico, Central America, Jamaica, Colombia to southeastern Brazil, Paraguay, Uruguay and northern Argentina.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with scalariform perforation plates; lateral pits alternate or opposite, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Prismatic calciumoxalate crystals abundant.

Leaves Alternate (distichous), simple, entire, with ? ptyxis. Stipules small, caducous; leaf sheath absent. Petiole vascular bundle transection D-shaped or deeply C-shaped; petiole also with wing bundles. Venation pinnate, brochidodromous. Stomata anomocytic (to anisocytic). Cuticular wax crystalloids? Mesophyll with calciumoxalate druses. Leaf margin usually serrate (rarely entire); salicoid teeth absent.

Inflorescences Axillary, racemose spike (Lozania) or catkin-like (Lacistema). Bracts large and imbricate (Lacistema), or small (Lozania).

Flowers Actinomorphic, small. Receptacle distinctly hollow, widened into fleshy concave disc. Hypogyny. Sepals (one or) two to six, unequal in size, free (sometimes absent). Petals absent. Nectariferous disc intrastaminal?, irregularly lobate, fleshy.

Androecium Stamen single. Filament inserted at or in disc, free from tepal. Anther basifixed, non-versatile, tetrasporangiate (thecae separated and sometimes individually stipitate), introrse, longicidal (dehiscing by longitudinal slits); connective widened. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of two or three connate carpels; median carpel adaxial. Ovary superior, unilocular. Style single, shortly bifid or trifid. Stigmas two or three, punctate, type? Pistillodium absent.

Ovules Placentation parietal. Ovules one or two per carpel (two to six per ovary), anatropous, pendulous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Funicle long, thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A usually single-seeded (sometimes two- or three-seeded) loculicidal? capsule, in Lozania on inner side with funicular? hair-like processes with thick unlignified cell walls.

Seeds Aril present? Testa sometimes carnose sarcotesta. Tegmen? Perisperm not developed. Endosperm copious, oily. Embryo usually long (rarely short), straight, well differentiated, chlorophyll? Cotyledons two, foliaceous. Germination?

Linales Bercht. et J. Presl, Přir. Rostlin: 239. Jan-Apr 1820 [‘Linicinae’]; Hugoniaceae Arn. in Wight et Arnott, Prodr. Fl. Ind. Orient. 1: 71. 22 Sep 1834

Fossils Uncertain. Fossil pollen grains assigned to Linaceae have been described from Miocene layers (Muller 1981).

Habit Bisexual, evergreen trees, shrubs, suffrutices or lianas, perennial, biennial or annual herbs. Some species are xerophytes. Hugonieae are often lianas with tendrils formed by basal branches of the inflorescences.

Vegetative anatomy Phellogen? Vessel elements with simple or scalariform perforation plates; lateral pits usually alternate, simple or bordered pits. Imperforate tracheary xylem elements long tough tenacious tracheids or fibre tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse?, or paratracheal aliform, lozenge-aliform, winged-aliform, or vasicentric (often difficult to define). Tyloses frequent. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace (i.a. Linum), or 3:3, trilacunar with three traces. Cortex in Hugonieae often with cristarque cells (in Lineae rarely). Prismatic calciumoxalate crystals often frequent.

Trichomes Hairs unicellular or multicellular, simple or branched, sometimes stellate; glandular hairs with multicellular head sometimes present.

Leaves Alternate (spiral) or opposite, simple, entire, usually with flat, involute or conduplicate ptyxis. Stipules lateral, small, caducous, or absent, in Linum modified into glands (extrafloral nectaries); leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata usually paracytic. Cuticular wax crystalloids as parallel platelets. Epidermis in Lineae often with mucilaginous idioblasts. Mesophyll at least in Hugonia with sclerenchymatous idioblasts. Leaf margin serrate or entire. Extrafloral nectaries sometimes (Linum) present on lamina.

Inflorescence Terminal or axillary, cymose of various shape (in Anisadenia spicate), often with partial inflorescences as cincinni or dichasia, or raceme, spike or panicle (in Hugonieae).

Flowers Usually actinomorphic (in Hugonieae often slightly zygomorphic). Pedicel articulated. Hypogyny. Sepals (four or) five, with imbricate quincuncial aestivation, persistent, free or connate at base. Petals (four or) five, with imbricate or contorted aestivation, caducous, free or connate at base (in Lineae usually clawed). Nectariferous glands extrastaminal, adnate to staminal tube or petal bases, or absent. Disc extrastaminal, annular or consisting of separate glands alternating with stamens. Flowers triheterostylous or diheterostylous in numerous species of Lineae, triheterostylous sometimes in Hugonieae (at least in Hugonia serrata).

Androecium Stamens in Lineae (four or) five, usually haplostemonous, antesepalous, alternipetalous (in Anisadenia obhaplostemonous, alternisepalous, antepetalous); in Hugonieae 5+5, diplostemonous. Filaments in Lineae often widened in lower part, connate at base into tube and sometimes with staminodia, free from tepals; filaments in Hugonieae connate into tube, free from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia two to five, extrastaminal, tooth-like or filiform or glandular, alternating with stamens, or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate or tricolporate (rarely tetracolpate, tetracolporate, hexacolpate, zonopolycolpate, zonopolyporate, zonopolyforate or inaperturate), starchy, shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate or acolumellate infratectum, reticulate or microreticulate, spinulate, gemmate or verrucate.

Gynoecium Pistil composed of (two or) three to five (to ten; in Anisadenia two) connate carpels; carpels antepetalous or median carpel adaxial. Ovary superior, unilocular or (bilocular or) trilocular to quinquelocular (to decemlocular; ovary sometimes unilocular only at apex); locules in Lineae usually (not in Anisadenia) divided into locelli by an individual incomplete secondary septum from carpellary midvein, synascidiate. Style single, simple, or stylodia (two or) three to five, free or connate at base. Stigmas capitate, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (in Hugonieae sometimes apical). Ovules (one or) two per carpel, anatropous, pendulous, epitropous? (micropyle directed upwards and outwards), bitegmic, usually crassinucellar (in Linum partially tenuinucellar). Micropyle bistomal or endostomal. Outer integument two or three cell layers thick. Inner integument three to twelve cell layers thick (Lineae). Obturator placental at least in Hugonieae. Parietal tissue four to six cell layers thick. Archespore sometimes multicellular. Endothelium present. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells in Linum three ephemeral nuclei (not cells). Endosperm development ab initio nuclear or helobial. Endosperm haustorium chalazal. Embryogenesis solanad.

Fruit In Lineae usually a septicidal capsule (rarely a drupe or nut; in Anisadenia a bipartite schizocarp); in Hugonieae usually a drupe (sometimes with septicidal dehiscence; sometimes schizocarp) with persistent calyx.

Seeds Aril absent; arillodium present in some Hugonieae. Seed coat exotegmic. Exotesta in Lineae with outer cell walls massively thickened. Mesotesta in Hugonieae with sclerotic cells. Endotesta in Hugonieae lignified. Tegmen highly multiplicative, sometimes reduced. Exotegmen fibrous. Endotegmen pigmented endothelium. Perisperm not developed. Endosperm copious to sparse or absent. Embryo usually straight (rarely somewhat curved), well differentiated, in Lineae oily and with chlorophyll. Cotyledons two. Germination phanerocotylar.

DNA Plastid genes rps16 and clpP and ORF244 absent (lost) (Linum grandiflorum). Mitochondrial coxI intron present (Linum).

Phytochemistry Insufficiently known. Ellagic and gallic acid present in at least Lineae. Saponins and cyanogenic compounds present in Hugonieae. Alkaloids? Proanthocyanidins? Flavonols not found.

Use Ornamental plants (Linum, Reinwardtia), textile and paper (fibres from Linum usitatissimum), seed oils (Linum usitatissimum, also as medicine for constipation treatment and as forage), timber, medicinal plants, fruits (Hugonia).

Habit Monoecious, usually climbing evergreen shrub (rarely tree). Tendrils consisting of modified leaves? (inflorescences?). Lateral bud present at branch bases.

Vegetative anatomy Phellogen subepidermal. Vessel elements with simple perforation plates; lateral pits? Imperforate tracheary xylem elements fibres with numerous minutely bordered pits. Wood rays uniseriate to multiseriate. Axial parenchyma scarce and paratracheal. Intraxylary phloem present. Phloem stratified. Sieve tube plastids S type? Nodes unilacunar? Crystals as solitary rhomboids and clusters.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules small; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids? Leaf margin serrate or crenate.

Inflorescence Axillary, panicle consisting of dense globular partial inflorescences.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with valvate aestivation, persistent, free or connate at base. Petals five, minute, with open aestivation, inflexed in bud, free. Nectariferous disc in female flowers as (staminodial?) glands, partially covering petal bases; cordate nectariferous glands in male flowers adnate to petals, in female flowers connate and forming quinquelobate nectariferous disc.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent (or as nectariferous lobes?).

Pollen grains Microsporogenesis simultaneous? Pollen grains tri- or tetracolporate, shed as monads, bicellular? at dispersal. Exine?, with ? infratectum, sculpturing?

Gynoecium Pistil composed of (four or) five connate antepetalous carpels. Ovary superior, synascidiate, (quadrilocular or) quinquelocular. Style single, (quadrilobate or) quinquelobate, very short or absent. Stigmas subulate, non-papillate, type? Pistillodium, very small, present in male flowers.

Ovules Placentation (apical to) axile. Ovules two per carpel, anatropous, pendulous, epitropous, bitegmic, weakly crassinucellar or incompletely tenuinucellar. Micropyle endostomal. Outer integument three cell-layers thick, without vascular bundles. Inner integument five or six cell-layers thick, without vascular bundles. Obturator funicular, small. Endothelium present. Megasporangium disintegrating. Nucellar beak absent. Nucellar cap absent. Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Seeds Aril present. Testa? Exotegmen fibrous. Endotegmen? Perisperm not developed. Endosperm copious. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Malesherbiales D. Don in C. F. P. von Martius, Consp. Regn. Veg.: 50. Sep-Oct 1835 [‘Malesherbiaceae’]

Distribution The Andes of southern Peru, northern Chile and western Argentina.

Habit Bisexual, evergreen shrubs or suffrutices, or usually perennial (rarely annual) herbs. Evil-smelling. Often densely hairy.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes cortical). Primary medullary strands narrow. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, scalariform or pseudoscalariform. Imperforate tracheary xylem elements libriform fibres with small pits (vascular tracheids sometimes present in late wood; also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal vasicentric, scanty, more or less scarce or absent. Tyloses sometimes present. Phloem fibres often present. Sieve tube plastids S type; sieve tubes with non-dispersive protein bodies? Nodes 1:1, unilacunar with one leaf trace, or 3:3, trilacunar with three traces. Rhomboidal calciumoxalate crystals single, few or absent; druses present in pits and cortex of some species.

Trichomes Hairs unicellular or multicellular, multiseriate, simple or branched; glandular hairs excreting nasty-smelling substance often present.

Leaves Alternate (spiral), usually simple (sometimes pinnately compound), entire or often deeply lobed, with ? ptyxis. Stipules absent (or foliaceous); small stipule-like leaves often present (prophylls of axillary buds or stipules?) at bases of leaves and bracts; leaf sheath absent. Stipules and/or prophylls and leaf primordia with colleters? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Leaf margin usually serrate or crenate (sometimes entire).

Inflorescence Axillary?, simple or compound panicle (rarely fascicle) or raceme, or flowers solitary.

Flowers Actinomorphic. Hypogyny. Sepals and petals connate into tubular, infundibuliform or campanulate persistent chartaceous 0,5–5 cm long hypanthium-like perigone. Sepals five, with valvate aestivation, free. Petals five, with valvate (to cochlear) aestivation, clawed, sometimes with thin corona with denticulate margin (as long as or slightly longer than remaining part of corolla), free. Androgynophore present, lobate and hairy, with nectary at base. Disc absent. Heterostyly present in almost all species.

Androecium Stamens five, protruding, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals, inserted at androgynophore. Anthers dorsifixed, versatile?, tetrasporangiate, introrse or extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tricolporate (rarely syncolporate), shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three (or four) connate carpels. Ovary superior, unilocular, inserted at androgynophore. Stylodia three (or four), filiform, free, inserted below ovary apex. Stigmas capitate or clavate, type? Pistillodium absent.

Ovules Placentation parietal. Ovules c. 30 to more than 100 per ovary, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument one to three cell layers thick. Chalazal part well developed and forming large outgrowth. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A stalked capsule, enclosed by persistent hypanthium-like perigonal tube.

Seeds Aril absent. Testa with pores. Exotestal cells arranged in rows? Endotesta well developed, with inner epidermis palisade of stone cells. Exotegmen palisade? Endotegmen persistent? Perisperm not developed. Endosperm fleshy, oily and with aleurone. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Phytochemistry Insufficiently known. Cyclopentenoid cyanogenic glycosides and/or cyclopentenylic fatty acids present. Tannins?

Malpighiopsida Bartl., Ord. Nat. Plant.: 227, 357. Sep 1830 [’Malpighinae’]; Malpighiineae Engl., Syllabus, ed. 2: 139. Mai 1898

Distribution Tropical and subtropical regions on both hemispheres, with their highest diversity in tropical South America.

Fossils Pollen grains have been described from the mid-Eocene and leaves, Banisteriophyllum and Malpighiastrum, are known from Early Cenozoic sites. Eoglandulosa warmanensis comprises flowers from the Eocene of southeastern North America and supposedly belong in Malpighiaceae.

Habit Usually bisexual (rarely polygamomonoecious), evergreen or deciduous trees, shrubs, suffrutices or lianas.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes deeply seated, almost near endodermis). Secondary lateral growth normal or anomalous. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal confluent, vasicentric, scalariform, reticulate, or banded, or absent. Tyloses often abundant. Secondary phloem often stratified into hard fibrous and soft parenchymatous zones. Intraxylary phloem present or absent. Sieve tube plastids S₀ type, without starch or protein inclusions. Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, unilacunar with one trace). Secretory cells abundant. Laticifers present in Galphimieae. Heartwood often with gum-like substances. Crystals absent?

Trichomes Hairs unicellular, furcate and often T-shaped – T-hairs, malpighiaceous hairs, balance hairs, medifixed hairs – or multicellular and multi-armed (sometimes simple or stellate); stinging hairs present in some genera (e.g. Malpighia); glandular hairs often abundant.

Leaves Usually opposite (rarely alternate [spiral, Acridocarpus] or verticillate), simple, usually entire (in Stigmaphyllon lobed), with ? ptyxis. Stipules cauline/interpetiolar (in e.g. Malpighia sometimes lobate), intrapetiolar (in trees and shrubs) or petiolar (i.a. Hiraea), free or connate (often rudimentary; in e.g. Acridocarpus absent); leaf sheath absent. Petiole vascular bundle transection arcuate; petiole and/or abaxial side of lamina often with usually two large flattened multicellular nectariferous glands. Venation usually pinnate (in Stigmaphyllon sometimes palmate). Stomata usually paracytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type). Epidermis with or without mucilaginous idioblasts. Domatia in Acridocarpus as hair tufts. Leaf margin usually entire (in Stigmaphyllon sometimes serrate); nectariferous glandular teeth rarely present on leaf margin.

Inflorescence Terminal or axillary, panicle or raceme- or umbel-like. Bracts often with extrafloral nectaries.

Flowers Actinomorphic or zygomorphic (often obliquely zygomorphic). Usually hypogyny (in Acridocarpus epigyny). Sepals four or five, with imbricate aestivation, free or slightly connate at base; usually with large paired abaxial oil-producing glands (epithelial elaiophores) at base and sticking to legs of neotropical oil-collecting bees. Petals five, with imbricate or contorted aestivation, often crumpled in bud, usually clawed and often with hairy, toothed or fringed margin, free; median petal adaxial or abaxial; one adaxial-lateral petal often larger and differently shaped and coloured than remaining petals. Nectariferous glands present at sepal bases in Old World species, or absent. Disc poorly developed or absent.

Androecium Stamens usually 5+5, obdiplostemonous (rarely two or 15 stamens or five antesepalous stamens, in one or three whorls). Filaments usually connate at base into tube, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (usually dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores); connective sometimes enlarged. Tapetum secretory, with multinucleate cells. Staminodia one to five, extrastaminal, or absent (antepetalous stamens sometimes staminodial or absent).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tri- to pentacolporate or tetra- to polyporate, shed as monads, bicellular at dispersal. Exine tectate to semitectate, with columellate or granular infratectum (sometimes with anastomosing elements), perforate, reticulate or microreticulate, often verrucate.

Gynoecium Pistil composed of usually three (sometimes four or five; in Acridocarpus two) usually connate carpels; median carpel usually obliquely orientated (carpels rarely almost free). Ovary usually superior (rarely inferior), usually trilocular (sometimes quadrilocular or quinquelocular; in Acridocarpus bilocular; rarely unilocular, apocarpous). Stylodia three (to five; in Acridocarpus two), usually free (style sometimes single, simple, subulate). Stigmas asymmetrically capitate or truncate, terminal or non-terminal, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile to apical. Ovule one per carpel, hemianatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle bistomal or endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Nucellar beak present. Megagametophyte usually tetrasporous, 16-nucleate, Penaea type (in Galphimia glabra disporous, Allium type). Endosperm development ab initio nuclear. Endosperm haustoria chalazal. Embryogenesis solanad (or adventitious). Nucellar polyembryony frequently occurring.

Fruit In syncarpous species a schizocarp (sometimes with carpophore, sometimes divided into two or three [to five] nut-, samara- or drupe-like mericarps), a nut or drupe, often with persistent (sometimes accrescent, sometimes wing-like) calyx and stamens; septicidal capsule present in someGalphimieae; fruiting carpels in some apocarpous species fusing and forming secondary syncarp.

Seeds Aril absent. Operculum? Seed coat usually exotegmic (sometimes endotegmic). Testa? Exotegmen in at least Byrsonima and Thryallis fibrous. Endotegmen often fibrous; endotegmic cells lignified, sometimes elongate. Perisperm not developed. Endosperm very sparse or absent. Embryo large, usually straight to somewhat curved (rarely hook-shaped, spirally twisted or circinate), oily, chlorophyll? Cotyledons two. Germination phanerocotylar or cryptocotylar.

DNA Plastid gene rps16 lost in at least Malpighia coccigera. Duplication of CYC genes. Mitochondrial coxI intron present (Malpighia).

Phytochemistry Insufficiently known. Condensed tannins present. Flavonols (kaempferol, quercetin), cyanidin, indole (harmidine) alkaloids, and triterpene saponins sometimes present. Iridoids? Cyanogenic compounds? Ellagic acid not found. Inulin sometimes present as carbohydrate reserve.

Use Ornamental plants, fruits (Malpighia), timber, narcotics (hallucinogens from Banisteriopsis and Diplopterys).

SystematicsMalphighiaceae are sister to Elatinaceae.

The clade [Galphimieae+[Acmanthereae+Byrsonimeae]] (Byrsonimoideae) is sister-group to the remaining Malpighiaceae (Davis & Anderson 2010).

Byrsonimoideae W. R. Anderson in Leandra 67(7): 6. Dec 1977 [1978]

10/215–240. Tropical America. Style subulate. Stigma terminal.

Byrsonimeae W. R. Anderson in Leandra 67(7): 7. Dec 1977 [1978]

3/150–175. Blepharandra (6; B. angustifolia, B. cachimbensis, B. fimbriata, B. heteropetala, B. hypoleuca, B. intermedia; southern Venezuela, Guyana, Amazonian Brazil), Diacidia (11; Colombia, Venezuela, Brazil, with their largest diversity in the Venezuelan Highlands), Byrsonima (135–160; southeastern Florida, southern Mexico, Central America, the West Indies, tropical South America to southeastern Brazil). – Tropical America.

Acmanthereae W. R. Anderson in Leandra 67(7): 11. Dec 1977 [1978]

3/c 23. Pterandra (c 15; Panamá to Brazil), Coleostachys (1; C. genipifolia; southern French Guiana, northern Brazil), Acmanthera (7; A. cowanii, A. duckei, A. fernandesii, A. latifolia, A. longifolia, A. minima, A. parviflora; Brazil). – Tropical America.

Galphimieae Nied. in Engler et Prantl, Nat. Pflanzenfam. III, 4: 53, 67. Dec 1890

4/41. Lophanthera (5; L. hammelii, L. lactescens, L. longifolia, L. pendula, L. spruceana; Costa Rica, Amazonian Brazil), Spachea (7; S. correae, S. elegans, S. herbert-smithii, S. martiana, S. membranacea, S. perforata, S. tricarpa; southern Central America, Cuba, Trinidad, northern tropical South America), Galphimia (27; southern Texas, Mexico to Nicaragua, tropical South America, with their highest diversity in Mexico), Verrucularia (2; V. glaucophylla, V. piresii; Brazil). – Tropical America. Laticifers articulated.

Malpighioideae Burnett, Outl. Bot.: 894, 1093, 1126. Jun 1835

c 63/1.170–1.205. Pollen grains tetraporate to polyporate. Stigma usually not terminal. Fruit winged. – The Acridocarpus clade is sister to the remaining Malpighioideae (Davis & Anderson 2010).

Acridocarpus clade

2/c 33. Acridocarpus (c 30; tropical and subtropical Africa, Madagascar, the Mascarene Islands, the Arabian Peninsula, India, one species, A. austrocaledonicus, in New Caledonia), Brachylophon (3; B. acuminatum, B. anastomosans, B. curtisii, tropical regions in the Old World). – Tropical regions in the Old World. Leaves spiral. Stipules absent. Epigyny. Two carpels fertile.

Mcvaughia clade

3/7. Mcvaughia (1; M. bahiana; Bahia in Brazil), Burdachia (3; B. duckei, B. prismatocarpa, B. sphaerocarpa; Amazonian Colombia and Venezuela, Guyana, Amazonian Brazil, eastern Peru), Glandonia (3; G. macrocarpa, G. prancei, G. williamsii; Amazonian Colombia, southern Venezuela, Amazonian Brazil). – Tropical South America.

Barnebya clade

1/2. Barnebya (2; B. dispar, B. harleyi; eastern Brazil). – Trees or lianas.

Ptilochaeta clade

4/13. Lasiocarpus (5; L. ferrugineus, L. multiflorus, L. ovalifolius, L. salicifolius, L. triflorus; Mexico), Ptilochaeta (6; P. bahiensis, P. densiflora, P. diodon, P. elegans, P. glabra, P. nudipes; tropical and subtropical South America), Dinemandra (1; D. ericoides; arid regions in Peru and Chile), Dinemagonum (1; D. gayanum; Chile). – Tropical and subtropical America.

Tristellateia clade

6/105–110. Echinopterys (2; E. eglandulosa, E. setosa; Mexico), Tristellateia (c 20; Madagascar, one species, T. africana, in tropical Africa, one species, T. australasiae, in tropical Asia and eastwards to tropical Australia and New Caledonia), Heladena (1; H. multiflora; southern Brazil, Paraguay, northeastern Argentina), Henleophytum (1; H. echinatum; Cuba), Thryallis (3; T. brachystachys, T. laburnum, T. parviflora; southern Brazil, Paraguay, Bolivia), Bunchosia (c 80; Mexico, Central America, the West Indies, tropical South America to southeastern Brazil and northern Argentina).

Hiraeeae Griseb. in C. F. P. von Martius, Fl. Bras. 12(1): 4, 75. 1 Jun 1858 [‘Hiraeaceae’]

5/77–82. Lophopterys (6; L. euryptera, L. floribunda, L. inpana, L. peruviana, L. splendens, L. surinamensis; tropical South America), Adelphia (4; A. hiraea, A. macrophylla, A. mirabilis, A. platyrachis; Central America, the West Indies, western South America), Hiraea (55–60; western Mexico, Central America, Grenada, St. Lucia, tropical South America to southeastern Brazil, Paraguay and northern Argentina), Psychopterys (8; southern Mexico, Belize, Guatemala), Excentradenia (4; E. adenophora, E. boliviana, E. primaeva, E. propinqua; northern South America). – Tropical America.

Tetrapterys clade

15/320–335. ‘Tetrapterys’ (c 90; tropical America; non-monophyletic), Niedenzuella (16; Central America, tropical South America), Dicella (7; D. bracteosa, D. nucifera, D. aciculifera, D. conwayi, D. julianii, D. macroptera, D. oliveirae; Costa Rica, tropical South America), Tricomaria (1; T. usillo; western Argentina), Carolus (6; C. anderssonii, C. chasei, C. chlorocarpus, C. dukei, C. renidens, C. sinemariensis; Mexico, Central America, the Lesser Antilles, tropical South America), Hiptage (30–35; Mauritius, Sri Lanka, northern Pakistan, southern Himalayas to southern China and Taiwan, Southeast Asia, Malesia to Fiji), Flabellariopsis (1; F. acuminata; tropical Africa),‘Heteropterys’ (130–140; Mexico, Central America, the West Indies, tropical South America southwards to southeastern Brazil and northern Argentina, one species, H. leona, in tropical West Africa; non-monophyletic), Christianella (5; C. glandulifera, C. mesoamericana, C. multiglandulosa, C. paludicola, C. surinamensis; Central America, tropical South America), Jubelina (6; J. grisebachiana, J. magnifica, J. riparia, J. rosea, J. uleana, J. wilburii; Central America, tropical South America), Alicia (2; A. anisopetala, A. macrodisca; tropical South America), Callaeum (11; western Texas, Mexico, Central America, tropical South America), Mezia (10; Panamá, tropical South America), Flabellaria (1; F. paniculata; tropical Africa), Malpighiodes (4; M. bracteosa, M. guianensis, M. leucanthele, M. liesneri; southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil).

Stigmaphyllon clade

15/445–450. Stigmaphyllon (113; the Ryukyu Islands, Taiwan, the Philippines to New Guinea, eastern Queensland, Solomon Islands, New Caledonia, Vanuatu, Micronesia, Palau, southern Mexico, Central America, the West Indies, tropical South America to northern Argentina), Bronwenia (11; southern Mexico, Central America, tropical South America), Diplopterys (c 45; Costa Rica, Panamá, tropical South America), ‘Banisteriopsis’ (90–95; southern Mexico, Central America, the West Indies, tropical South America; non-monophyletic), ‘Sphedamnocarpus’ (18; tropical and southern Africa, Madagascar; non-monophyletic; incl. Philgamia?), Philgamia (4; P. brachystemon, P. denticulata, P. glabrifolia, P. hibbertioides; Madagascar; in Sphedamnocarpus?), Peixotoa (c 30; Brazil, Paraguay, Bolivia), Gallardoa (1; G. fischeri; Argentina), Mionandra (1; M. camareoides; Bolivia, Paraguay, Argentina), Cordobia (2; C. argentea, C. fischeri; South America), Cottsia (3; C. californica, C. gracilis, C. linearis; southern United States, northern Mexico), ’Janusia’ (18; California to Argentina; paraphyletic), ’Aspicarpa’ (c 40; southern United States, Mexico, Central America, the West Indies, tropical and subtropical South America; paraphyletic; incl. Gaudichaudia?), ‘Gaudichaudia’ (c 60; Mexico to Bolivia; polyphyletic; in Aspicarpa?), Camarea (9; eastern South America). – Pantropical, southern United States.

Ectopopterys clade

1/1. Ectopopterys (1; E. soejartoi; Colombia, Peru). – Liana.

Amorimia clade

1/10. Amorimia (10; tropical South America). – Lianas.

Malpighieae DC., Prodr. 1: 577. Jan (med.) 1824

c 10/155–170. ’Mascagnia’ pro parte (35–50; Mexico to Argentina; non-monophyletic), Calcicola (2; C. parvifolia, C. sericea; Mexico), ‘Malpighia’ (c 45; Florida, Mexico, Central America, the West Indies, tropical South America; non-monophyletic), Aspidopterys (c 25; tropical Asia), Caucanthus (3; C. albidus, C. auriculatus, C. edulis; eastern and northeastern Africa, the Arabian Peninsula), Triaspis (c 15; tropical and southern Africa), Digoniopterys (1; D. microphylla; Madagascar), Rhynchophora (2; R. humbertii, R. phillipsonii; Madagascar), Madagasikaria (1; M. andersonii; southern Madagascar), Microsteira (c 25; Madagascar). – Eastern Africa, Madagascar, the Arabian Peninsula, tropical Asia, tropical America. The endemic Malagasy genera Madagasikaria, Microsteira and Rhynchophora form a well supported clade, according to Davis (2002).

Unplaced Malpighiaceae

Rudolphia (1; R. planisiliqua; Bolivia).

Cladogram (simplified) of Malpighiaceae based on DNA sequence data (Davis & Anderson 2010).

Habit Andromonoecious, evergreen tree. Bark fibrous (resembling Juniperus bark). Flowers evil-smelling. Buds perulate.

Vegetative anatomy Phellogen ab initio subepidermal. Secondary lateral growth anomalous. Vessel elements with simple perforation plates; lateral pits opposite or alternate, bordered pits. Imperforate tracheary xylem elements tracheids or libriform fibres with small bordered pits, non-septate? Wood rays multiseriate?, heterocellular. Axial parenchyma apotracheal diffuse. Secondary phloem stratified into concentric rings of alternating sieve tissue and sclerenchyma. Sieve tube plastids S type. Nodes 5:5, pentalacunar with five leaf traces, and two phloic vascular bundles. Secretory cavities absent. Cristarque cells absent. Calciumoxalate druses present.

Leaves Opposite, simple, entire, coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Paired colleters present along leaf margin and in leaf axil outside axillary bud. Petiole vascular bundle transection arcuate; petiole with numerous bundles, variously orientated. Venation pinnate; tertiary venation highly reticulate. Stomata anomocytic. Cuticular waxes absent. Mesophyll with mucilaginous idioblasts, without sclerenchymatous idioblasts. Calciumoxalate druses present adjacent to vascular strands of midvein. Leaf margin crenate.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with imbricate aestivation, persistent, connate at base. Petals five, with imbricate aestivation, recurved, caducous, free. Nectary absent. Disc absent.

Androecium Stamens c. 50 to more than 100, spirally arranged, arising from five vascular bundles. Filaments thin, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains (2–)3(–4)-porate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, finely striate.

Gynoecium Pistil composed of 16 to 19 (to 25) connate carpels, inserted on and adnate to central column; ventral sutures open below ovular points of insertion. Ovary superior, 16- to 25-locular. Stylodia 16 to 25, free, marginal, inserted in characteristic way on outer carpel margins (cf. the name Medusagyne). Stigmas capitate, discoid, type? Pistillodium absent.

Ovules Placentation axile. Ovules two to five per carpel, anatropous, upper ovules ascending to horizontal and epitropous, lower ovules descending to pendulous and apotropous, bitegmic, weakly crassinucellar. Micropyle ?-stomal. Outer integument three or four cell layers thick. Inner integument three or four cell layers thick. Funicle long. ‘False endothelium’ present on surface of megasporangium. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit An assemblage of ridged, verrucose, septicidal follicles with persistent central columella, stigmas and calyx and carpels acropetally detached – valves detaching from central columella from base upwards except at apex, eventually umbrella-like with ventricidal carpels –, adaxially dehiscing (fruit anatomy similar to Caryocaraceae), also interpreted as a schizocarp with 16 to 25 follicle-like mericarps. Exocarp without lacunae.

Seeds Aril absent. Testa winged. Exotesta somewhat thickened. Endotesta crushed. Exotegmen fibrous. Endotegmen crushed. Perisperm not developed. Endosperm thin. Embryo straight, chlorophyll? Cotyledons two. Germination phanerocotylar, epigeal or hypogeal.

Phytochemistry Insufficiently known. Tannins and phenolic compounds present.

Ochnales DC. ex Bercht. et J. Presl, Přir. Rostlin: 224. Jan-Apr 1820 [‘Ochnaceae’]; Sauvagesiaceae (Ging. ex DC.) Dumort., Anal. Fam. Plant.: 44, 49. 1829; Lophiraceae Loudon, Hort. Brit.: 513. 30 Aug 1830 [‘Lophireae’]; Sauvagesiales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 50. Sep-Oct 1835 [‘Sauvagesiaceae’]; Gomphiaceae DC. ex Schnizlein, Iconogr. Fam. Regni Veg. 4: ad t. 248. 1843-1870; Luxemburgiaceae Soler., Syst. Anat. Dicot. Ergänz.: 77, 79. Mai 1908; Ochnanae Doweld, Tent. Syst. Plant. Vasc.: xxxiii. 23 Dec 2001

Vegetative anatomy Phellogen ab initio superficial. Cortical and medullary vascular bundles usually present. Primary vascular tissue cylinder. Secondary lateral growth normal. Vessel elements usually with simple (sometimes also scalariform) perforation plates; lateral pits alternate, usually bordered pits. Vestured pits often present (absent in Sauvagesia). Imperforate tracheary xylem elements usually fibre tracheids (sometimes libriform fibres) with simple or bordered pits, usually septate (occasionally non-septate; vasicentric tracheids rare). Wood rays uniseriate or multiseriate, usually heterocellular (rarely homocellular). Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, vasicentric, or unilateral, or metatracheal. Phloem sometimes (i.a. in Godoya) stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids S type. Nodes ≥3:≥3, trilacunar or multilacunar with three or more? leaf traces. Mucilage cells and mucilage ducts often present. Cortex often with cristarque cells. Hyaline latex present in some species. Heartwood often with gum-like substances. Silica bodies present in some species. Prismatic calciumoxalate crystals often abundant.

Trichomes Hairs usually absent (simple hairs sometimes present); glandular hairs sometimes present.

Leaves Alternate (spiral or distichous), usually simple (rarely pinnately compound), entire, often coriaceous, usually with conduplicate to flat ptyxis. Stipules extrapetiolar, scale-like, often fimbriate, in Ouratea with extrafloral nectaries; leaf sheath absent. Petiole vascular bundle transection annular (sometimes several bundles arcuately arranged); petiole sometimes with medullary bundles. Venation pinnate, brochidodromous; secondary veins stout and often densely parallel and/or tertiary veins parallel. Stomata usually paracytic (rarely anomocytic). Cuticular waxes absent? Epidermis with or without mucilaginous idioblasts. Mesophyll often with sclerenchymatous idioblasts, forming continuous subepidermal layer on adaxial side. Cristarque cells often abundant. Leaf margin serrate, crenate or entire.

Inflorescence Terminal or axillary, panicle, fascicle, raceme or umbel, or flowers solitary.

Flowers Usually actinomorphic (rarely somewhat zygomorphic due to non-uniformly developed androecium). Pedicel articulated. Hypogyny. Sepals (three to) five (to ten), with imbricate aestivation, almost membranous (in Lophira non-uniformly growing), sometimes persistent, usually whorled (rarely spiral), free or connate at base. Petals (four or) five (to ten), usually with contorted (rarely imbricate) aestivation, sometimes clawed, caducous, usually whorled (rarely spiral), free. Nectary absent. Disc absent.

Androecium Stamens five to 25 (to more than 300), in one to five whorls, often in five groups, sometimes on androphore, sometimes centrifugally developing. Filaments free or more or less connate, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, often latrorse, usually poricidal (dehiscing by apical pores; sometimes longicidal, dehiscing by longitudinal slits; sometimes locellate). Tapetum secretory. Staminodia in Sauvagesioideae (five or) ten to c. 25, in one or several whorls, sometimes petaloid (sometimes connate into tubular corona), or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate, shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine tectate, columellate infratectum, microperforate or striate-rugulate.

Gynoecium Pistil composed of (one to) five to ten (to 15) carpels free in upper part and connate in lower part (in Ouratea secondarily free), or carpels paracarp and entirely connate (antesepalous when three median carpels adaxial). Ovary superior, usually unilocular to quinquelocular (rarely up to 15-locular by placental intrusions), often deeply lobate. Style single, simple, usually long (sometimes short), sometimes hollow, sometimes gynobasic (e.g. in Ouratea). Stigmas punctate or somewhat lobate, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation axile, parietal or intrusively parietal (marginal?). Ovules one, two or five to more than 50 per carpel, anatropous to campylotropous, usually ascending (rarely pendulous), apotropous or epitropous, usually bitegmic, usually more or less connate (in Lophira unitegmic), incompletely tenuinucellar or weakly crassinucellar. Micropyle bistomal or endostomal (or integuments connate), often zigzag. Outer integument two to four cell layers thick. Inner integument two to four cell layers thick. Hypostase often present. Endothelium absent? Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Antipodal cells usually persistent, often enlarged. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis? Polyembryony may occur.

Fruit A berry, drupe (sometimes an assemblage of drupes) or septicidal capsule (in Ouratea schizocarp with three to ten drupaceous mericarps situated on fleshy carpophore; in Lophira capsule with persistent calyx with two wing-like sepals).

Seeds Aril absent. Testa well developed or reduced and thin (Lophira), often winged, often vascularized. Endotesta with small crystalliferous cells. Exotegmen often fibrous. Endotegmen? Perisperm not developed. Endosperm sparse or absent, with or without oils. Embryo usually straight (sometimes curved), chlorophyll? Cotyledons two. Germination phanerocotylar or cryptocotylar.

Phytochemistry Biflavonyls/biflavonoids, C-glycoflavones, flavones, biflavones, lophirone, Ouratea-catechins, cyanidin,vismiones, and anthrones (3-O-geranylemodin anthrone etc.) present. Condensed tannins present. Triterpenes and alkaloids rare. Flavonols, ellagic acid, saponins, and cyanogenic compounds not found.

Use Ornamental plants, medicinal plants, timber, roofing and basketing (Cespedesia).

Systematics Ochnaceae belong in a trichotomy also comprising Medusagyne (Medusagynaceae) and Quiinaceae, although in some analyses Medusagyne is sister to Quiinaceae.

A plausible topology of the Ochnaceae phylogeny is the following, according to Schneider & al. (2014):

[Testulea+[Luxemburgioideae+[Ochnoideae+Sauvagesioideae]]]

Testulea clade

1/1. Testulea (1; T. gabonensis; Gabon). – Testulea is sister-group to the remaining Ochnaceae, according to Schneider & al. (2014).

[Luxemburgioideae+[Ochnoideae+Sauvagesioideae]]

Luxemburgioideae Planch. ex Endl., Gen. Plant. Suppl. 5: 98. 1850 [‘Luxembergieae’]

2/22. Luxemburgia (18; Brazil), Philacra (4; P. auriculata, P. duidae, P. longifolia, P. steyermarkii; Venezuela, northern Brazil). – Venezuela, Brazil. Androecium and gynoecium obliquely zygomorphic in bud. One adaxial stamen fertile; remaining (staminodial) stamens grouped on one side of gynoecium. Filament connate. Anthers sometimes connate. Exine with small perforations. Pistil composed of three connate carpels. n = ? – Luxemburgioideae seem to be sister-group to the remaining Ochnaceae.

[Ochnoideae+Sauvagesioideae]

Staminodia separate, forming lobate disc or corolla-like tube, or absent. Exine striate-rugulate.

Ochnoideae Burnett, Outlines Bot.: 886, 1093, 1125. Feb 1835 [‘Ochnidae’]

9/c 385. Lophira (2; L. alata, L. lanceolata; tropical West and Central Africa), Elvasia (14–15; Central America, tropical South America), Campylospermum (c 65; tropical central and East Africa, Madagascar, tropical Asia), Idertia (2; I. axillaris, I. morsonii; western and central tropical Africa, São Tomé), Brackenridgea (10; tropical regions in the Old World), Ouratea (c 200; tropical regions on both hemispheres), Rhabdophyllum (4; R. letestui, R. reflexum, R. ‘rigidum’, R. thonneri; western and central tropical Africa), Ochna (c 85; tropical regions in the Old World). – Unplaced Ochnoideae Perissocarpa (3; P. ondox, P. steyermarkii, P. umbellifera; Venezuela, Peru, northern Brazil). – Tropical, with their largest diversity in Brazil. Leaves distichous. Vessel and parenchyma pits not unilaterally compound. Stipules in Ouratea semi-intrapetiolar. Petiole sometimes with inverted medullary vascular bundle and subepidermal fibres. Sepals almost membranous (in Lophira non-uniformly growing). Petals with contorted aestivation. Stamens diplostemonous or obdiplostemonous, centripetally developing. Anthers sometimes longicidal (dehiscing by longitudinal slits). Pistil composed of (two to) five (to 15; in Lophira two) connate carpels, sometimes present on short gynophore. Style sometimes gynobasic. Floral receptacle sometimes widened. Ovules usually one (in Lophira numerous) per locule, apotropous. Integuments connate (except occasionally at apex), together seven to 17 cell layers thick. Outer integument in Ochna three or four cell layers thick. Inner integument in Ochna two or three cell layers thick. Hypostase usually present. Fruit usually a drupe, with persistent stamens (in Lophira with two sepals modified into wings). Testa vascularized, without layer of small crystalliferous cells. Fibrous exotegmen absent. Endosperm absent. Cotyledons massive, variously arranged. x = 12–14.

Sauvagesioideae Beilschm. in Flora 16(Beibl. 7): 88, 110. 14 Jun 1833 [‘Sauvagesieae’]

16/90–95. Blastemanthus (2; B. gemmiflorus, B. grandifloras; northeastern South America), Godoya (2; G. antioquiensis, G. obovata; the Andes in Colombia, Peru and Bolivia), Rhytidanthera (2; R. magnifica, R. splendida; Colombia, Venezuela), Cespedesia (1–6; C. spathulata; Central America, tropical South America), Krukoviella (1; K. disticha; Peru, Brazil), Fleurydora (1; F. felicis; Guinea), Poecilandra (2; P. pumila, P. retusa; northern tropical South America), Wallacea (3; W. insignis, W. multiflora, W. riparia; northern South America), Neckia (1; N. serrata; West Malesia, the Philippines), Schuurmansia (18; Central Malesia to New Guinea), Schuurmansiella (1; S. angustifolia; northwestern Borneo), Euthemis (2; E. leucocarpa, E. minor; Southeast Asia to Borneo), Tyleria (14; Venezuela, the Guayana Highlands), Adenarake (2; A. macrocarpa, A. muriculata; southern Venezuela, northern BrazilVenezuela), Sauvagesia (c 40; tropical regions on both hemispheres, with their highest diversity in tropical South America). – Unplaced Sauvagesioidae Indosinia (1; I. involucrata; southern Vietnam). – Pantropical, with their highest diversity in tropical South America. Rarely perennial herbs. Vestured pits absent in Sauvagesia. Leaves spiral, often with conduplicate-flat ptyxis (in Rhytidanthera compound). Flowers rarely zygomorphic (with late developing zygomorphy comprising androecium and gynoecium). Outer sepals sometimes smaller than remainder. Sauvagesia with five petaloid antepetalous staminodia and five antesepalous stamens. Exine sometimes with small perforations. Pistil composed of two, three or five connate carpels; when three, then median carpel adaxial. Outer integument approx. two cell layers thick. Inner integument three or four cell layers thick. Testa often winged. Exotestal cells large, detached. Endotestal cells crystalliferous. Endosperm with aleurone. x = 18. – Blastemanthus may be sister to the remaining Sauvagesioideae.

Cladogram (simplified) of Ochnaceae based on Schneider & al. (2014).

Engler et Gilg in Engler’s Syllabus, ed. 7: 223. Oct 1912-Mar 1913, nom. cons.

Bennettiaceae R. Br. ex Schnizlein, Iconogr. Fam. Regni Veg. 3: ad t. 172**. 1846-1866 [’Bennettieae’], nom. illeg.; Pandales Engl. et Gilg in Engler, Syllabus, ed. 7: 223. Oct 1912-Mar 1913

Habit Dioecious, evergreen trees or shrubs. Buds arising in axils of leafy short shoots, although often not in true leaf axils.

Vegetative anatomy Phellogen? Primary medullary strands alternately wide and narrow. Vessel elements usually with scalariform (sometimes also simple) perforation plates; lateral pits alternate or opposite, simple pits. Non-vestured pits present (Microdesmis). Imperforate tracheary xylem elements tracheids or fibre tracheids with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, scalariform, or banded. Tyloses frequent. Sieve tube plastids S type?; sieve tubes with non-dispersive protein bodies. Nodes? Latex and laticifers absent. Cortex with cristarque cells. Prismatic calciumoxalate crystals abundant.

Trichomes Hairs unicellular or multicellular, uniseriate; glandular hairs sometimes present on floral parts.

Leaves Alternate (spiral and reduced on orthotropic shoots; distichous on plagiotropic short shoots resembling pinnately compound leaves, especially in Galearia and Panda), simple, entire or pinnately lobed, with involute ptyxis. Stipules small (asymmetrical in Panda), usually persistent (caducous in Panda); leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate; one vein proceeding into persistent transparent apex of leaf tooth. Stomata anomocytic (Microdesmis), paracytic (Galearia) or cyclocytic (Panda). Cuticular waxes absent. Lamina with or without glandular dots (pellucid-punctate in Microdesmis). Mesophyll with sclerenchymatous idioblasts containing calciumoxalate as druses or solitary prismatic crystals. Leaf margin serrate or entire.

Inflorescence: Terminal or axillary, fascicle, thyrsoid or panicle, or flowers solitary axillary.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with imbricate or open (Panda) aestivation, free or connate (sepals of female flowers in Microdesmis usually persistent, with basal glandular hairs). Petals five, with imbricate, contorted or valvate aestivation, free (in Galearia each enclosing one or two stamens). Nectary absent. Disc very small or absent.

Androecium Stamens five antesepalous, or ten, 15 or 5+5 in one or two whorls. Filaments usually free from each other, free from or partially adnate to tepals. Anthers basifixed, non-versatile, tetrasporangiate, usually introrse (sometimes latrorse), longicidal (dehiscing by longitudinal slits); connective sometimes prolonged. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate or punctate.

Gynoecium Pistil composed of two to five connate carpels. Ovary superior, bilocular to quinquelocular. Style single, simple, or stylodia two to five, free or connate at base (rarely absent). Stigmas not widened, capitate, stigma sometimes bilobate, type? Male flowers with pistillodium.

Ovules Placentation axile-apical. Ovule usually one per carpel, orthotropous (Panda) or anatropous, pendulous, epitropous, bitegmic, weakly crassinucellar. Micropyle endostomal. Outer integument three to five cell layers thick. Inner integument three to five cell layers thick. Obturator absent. Megagametophyte monosporous, Polygonum type. Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Carunculus absent. Seed coat exotegmic. Exotesta possibly absent. Endotesta possibly absent. Exotegmen tracheoidal, fibrous. Endotegmen tanniniferous. Perisperm not developed. Endosperm copious, oily. Embryo well differentiated, chlorophyll? Cotyledons two, thin and flat, oily. Germination phanerocotylar.

Phytochemistry Very insufficiently known. Alkaloids and/or saponins sometimes present.

Passiflorales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 237. Jan-Apr 1820 [‘Passifloreae’]; Paropsiaceae Dumort., Anal. Fam. Plant.: 37, 42. 1829; Smeathmanniaceae Mart. ex Perleb, Clav. Class.: 33. Jan-Mar 1838 [‘Smeathmannieae’]; Passifloropsida Brongn., Enum. Plant. Mus. Paris: xxix, 108. 12 Aug 1843 [’Passiflorineae’]; Modeccaceae Horan., Char. Ess. Fam: 146. 30 Jun 1847 [‘Turneraceae s. Modeccaceae’]; Passiflorineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 462. 15 Nov 1895

Distribution Tropical, subtropical and warm-temperate regions on both hemispheres, with their highest diversity in tropical Africa and tropical America.

Fossils Seeds assigned to Passiflora have been described from the Miocene of Europe.

Habit Usually bisexual (in, e.g., Adenia dioecious), usually perennial or annual herbs, usually climbing and twining with tendrils, or evergreen shrubs or lianas (rarely small trees). Tendrils simple or branched, axillary, usually consisting of modified pedicels or inflorescences, often oblique relative to the branch. Some species are xerophytes.

Vegetative anatomy Phellogen? Passiflora and its closely allied relatives possessing stem collenchyma. Axillary buds usually superposed, with accessory bud originating above axillary inflorescence/tendril bud. Secondary lateral growth usually normal (in Adesmia anomalous, from concentric cambia). Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with bordered pits, non-septate (also vasicentric tracheids). Wood rays multiseriate?, homocellular or heterocellular. Axial parenchyma usually apotracheal diffuse? Wood often fluorescent. Intraxylary phloem present in some lianas. Sieve tube plastids S type; sieve tubes with non-dispersive protein bodies? Nodes 3:3, trilacunar with three leaf traces. Calciumoxalate as druses and solitary prismatic crystals.

Trichomes Hairs unicellular or multicellular, usually simple (sometimes stellate).

Leaves Alternate (spiral), usually simple (rarely palmately compound), entire or palmately lobed, with conduplicate ptyxis (sometimes modified conduplicate, V-shaped and at end of each arm Λ-shaped). Stipules often small and caducous, sometimes foliaceous (absent in Androsiphonia and Barteria); leaf sheath absent. Stipules and/or prophylls and leaf primordia with colleters? Petiole and/or stipules often with extrafloral nectaries without nectarostomata. Petiole vascular bundle transection? Venation pinnate, pedate or palmate; vein proceeding into transparent and caducous tooth apex? Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids as rosettes of platelets (Fabales type). Mesophyll with or without sclerenchymatous idioblasts. Epidermis sometimes with mucilaginous idioblasts. Calciumoxalate druses and solitary prismatic crystals frequent. Leaf margin usually serrate (sometimes entire). Lamina sometimes with extrafloral nectaries.

Inflorescence Usually axillary, racemose or cymose (flowers rarely solitary). Flowers usually subtended by three bracts.

Flowers Usually actinomorphic (rarely zygomorphic, e.g. Passiflora mucronata), usually large. Hypanthium usually absent (present in some species of Adenia). Usually half epigyny (rarely hypogyny). Sepals (three to) five (to eight), with imbricate aestivation, often petaloid, persistent, often connate at base, often adnate to petal bases. Petals (three to) five (to eight; rarely absent), with imbricate aestivation, usually with staminodial corona at base consisting of one to several whorls of c. 15 to more than 50 extrastaminal scales, hairs or filiform lobes, usually free (rarely connate at base). Sepals and petals often adnate to each other at base, sometimes forming common perianth tube. Annular nectariferous disc or five nectaries, hypogynous, staminodial, extrastaminal or alternating with stamens, inserted on perianth tube, or absent. Receptacle often distinctly hollow, usually with androphore; androgynophore usually present (in, e.g., Adenia gynophore).

Androecium Stamens usually five or eight, antesepalous, alternipetalous (sometimes four or c. 20 to more than 60). Filaments usually free (in, e.g., Adenia sometimes connate at base; in Androsiphonia connate into tube surrounding gynophore), sometimes adnate to and forming tube surrounding gynoecium. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia numerous (c. 15 to more than 50), extrastaminal to intrastaminal, often petaloid, as showy corona or disc.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3–12-colporate to 3–12-colpoidorate (rarely hexaporate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, sometimes echinate.

Gynoecium Pistil composed of (two or) three (to seven) connate carpels. Ovary semi-inferior (rarely superior), unilocular, stipitate. Stylodia (two or) three (to seven), free or partially connate (rarely entirely connate into single, simple style). Stigmas capitate to discoid (in Adenia split), sometimes lobate, usually with multicellular (in Adenia unicellular) papillae or non-papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal. Ovules usually numerous (in Dilkea few) per ovary, usually anatropous (rarely orthotropous), bitegmic, crassinucellar. Funicle often long. Micropyle bistomal, sometimes Z-shaped (zig-zag). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megasporangium often pointed at apex. Megagametophyte monosporous, Polygonum type. Synergids sometimes of different shape, sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually onagrad (rarely piperad).

Seeds Seeds with arilloid, often flattened. Testa multiplicative, hard, often hairy (rarely winged). Exotesta often sarcoexotesta. Endotestal cells large, with crystals and often lignified walls. Exotegmen palisade. Endotegmen persistent? Perisperm not developed. Endosperm copious, ruminate (Passiflora foetida) or non-ruminate, oily and proteinaceous. Embryo straight, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

DNA Plastid ORF2280 absent (lost). Plastid gene rps16 lost in Passiflora. Plastid gene rpoC1 in Passiflora without intron. Plastid gene rpl22 in Passiflora probably transferred from plastid genome to nuclear genome.

Phytochemistry Flavonols, flavone-C-glycosides, ellagic and gallic acids, alkaloids, valine- and isoleucine-derived cyanogenic glycosides, cyclopentenoid cyanogenic glycosides and/or cyclopentenyl fatty acids (from the gynocardic group), saponins, and cinnamic acid derivatives present. Tannins and proanthocyanidins not found.

Klotsch in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1859: 241, 246. 10-30 Mar 1859

Habit Usually dioecious (sometimes monoecious, especially in Pera), evergreen trees or shrubs (sometimes perennial herbs, more or less lignified at base).

Vegetative anatomy Phellogen? Cortical and medullary vascular bundles absent. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays uniseriate or multiseriate (in Pera uniseriate), homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal reticulate, vasicentric, or banded, abundant. Tyloses often frequent (sometimes sclerotic). Intraxylary phloem present at least in Pera. Sieve tube plastids S type? Nodes? Secretory cavities absent. Wood in some species with lysigenous radial ducts. Laticifers present in some species. Heartwood sometimes with gum-like substances. Prismatic calciumoxalate crystals abundant.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple (in Pera usually stellate or lepidote; inflorescences in Pogonophora sometimes with malpighiaceous hairs), or absent.

Leaves Usually alternate (spiral or distichous; rarely opposite), simple, entire, with asymmetrical leaf base, often coriaceous, with ? ptyxis. Stipules usually small (in Chaetocarpus and Trigonopleura large), often caducous, or absent; leaf sheath absent. Petiole vascular bundle transection interrupted arcuate to annular or complete annular; petiole sometimes with central plate and wing bundles. Venation pinnate; veins sometimes proceeding into transparent caducous teeth or marginal spines. Stomata? Cuticular wax crystalloids? Epidermis with mucilaginous idioblasts. Lamina without secretory cavities, sometimes pellucid-punctate. Leaf margin entire to prickly dentate.

Inflorescence Axillary, panicle, fascicle, thyrsoid or capitate glomerulus (female flowers sometimes solitary); male flowers in Pera often surrounded by reduced sterile female flowers. Bracts one or two small outer, free, and two larger involucrate inner, more or less connate.

Flowers Actinomorphic, small? Hypogyny. Sepals in male flowers (two to) four or five (to seven), with imbricate aestivation, free or connate (in Pera sometimes absent; sometimes absent in female flowers). Petals usually four or five, with imbricate aestivation, sometimes unguiculate (absent in Pera and Chaetocarpus; sometimes absent in female flowers), in Pogonophora with rigid hairs on adaxial side. Nectary? Disc usually consisting of five to numerous usually intrastaminal glands (nectariferous glands?) in one to three whorls (disc in Chaetocarpus extrastaminal, entire to somewhat lobate [crenellate]; disc absent in Pera).

Androecium Stamens (two to) five to eight (to 20). Filaments connate into tube at least in lower part, usually free from tepals. Anthers usually basifixed (sometimes dorsifixed), non-versatile?, tetrasporangiate, introrse or extrorse (sometimes latrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers with or without staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate (rarely intectate), with columellate infratectum, perforate or punctate to micropunctate-rugulate, sometimes psilate.

Gynoecium Pistil composed of three (to nine) connate carpels or one carpel. Ovary superior, trilocular (to novemlocular) or unilocular; ovary septa membranous and not distinctly vascularized. Style single, simple or trilobate (lobes often bifid), or absent. Stigmas one or three, papillate, peltate to fimbriate (sometimes valve-shaped?), type? Male flowers with or without pistillodium (entire, bilobate or trilobate).

Ovules Placentation apical. Ovules one per carpel, anatropous?, pendulous, epitropous, bistomal, crassinucellar. Micropyle usually endostomal (rarely exostomal). Outer integument three to six cell layers thick. Inner integument three to six cell layers thick. Obturator placental. Archespore bicellular or tricellular. Megasporangium approx. two cell layers thick, early degenerating. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A single-seeded drupe or drupaceous usually septicidal capsule with fleshy or spongy mesocarp and lignified endocarp, often with persistent tepals and stigmas; septicidal capsule usually dehiscing into three mericarps, each partially dehiscing into two valves (capsule rarely loculicidal and dehiscing into six valves). Fruit septa membranous and not distinctly vascularized. Capsular valves dehiscing along midvein and at base often remaining adnate to fruit stalk, central columella (usually thin, in Trigonopleura three-winged) splitting longitudinally into usually three parts.

Seeds Carunculus (in Trigonopleura also aril) present. Seed coat exotestal-exotegmic. Testa vascularized. Exotesta palisade, lignified, usually tanniniferous. Exomesotesta sclereidal. Endotestal cells with calciumcarbonate? Exotegmen usually tracheoidal (in Pogonophora palisade), sometimes U-shaped in cross-section; exotegmic cells cuboid? Endotegmen absent. Perisperm not developed. Endosperm usually copious, fleshy (rarely absent). Embryo straight, well differentiated, chlorophyll? Cotyledons two, wide and flat. Germination?

Stilaginaceae C. Agardh, Aphor. Bot.: 199. 13 Jun 1824 [‘Stilagineae’]; Antidesmataceae Loud., Hort. Brit.: 534. 30 Aug 1830 [’Antidesmeae’]; Stilaginales C. Agardh in C. F. P. von Martius, Consp. Regn. Veg.: 14. Sep-Oct 1835 [‘Stilagineae’]; Scepaceae Lindl., Intr. Nat. Syst. Bot., ed. 2: 171. 13 Jun 1836; Aporosaceae Lindl. ex Planch. in Ann. Sci. Nat., Bot. Ser. 4, 2: 265. 1854 [’Aporoseae’]; Porantheraceae (Pax) Hurus. in J. Fac. Sci. Univ. Tokyo, ser. III, 6: 224. 15 Aug 1954; Bischofiaceae (Müll. Arg.) Airy Shaw in Kew Bull. 18: 252. 8 Dec 1965; Hymenocardiaceae Airy Shaw in Kew Bull. 18: 261. 8 Dec 1965; Uapacaceae (Müll. Arg.) Airy Shaw in Kew Bull. 18: 270. 8 Dec 1965; Phyllanthales Doweld, Tent. Syst. Plant. Vasc.: xxxiii. 23 Dec 2001

Distribution Tropical and subtropical regions on both hemispheres, with few species in warm-temperate regions.

Fossils Uncertain. Paraphyllanthoxylon, fossil wood from Cretaceous layers of South Africa and North America, may represent Phyllanthaceae or Euphorbiaceae. A fossil flower attributed to Antidesma has been found in Baltic amber (Willemstein 1987).

Habit Usually monoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees or shrubs (sometimes climbing), perennial or annual herbs. Some species are xeromorphic shrubs or herbs with ericoid leaves (some species of Phyllanthus have phyllocladia; one species of Phyllanthus is limnic and floating). Branches often dimorphic.

Vegetative anatomy Ectomycorrhiza present in Uapaca. Phellogen ab initio superficial? Primary medullary strands narrow (Hymenocardia). Vessel elements with simple or scalariform perforation plates; lateral pits alternate, simple and/or bordered pits. Vestured pits present in Bridelia and allies. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate (in ‘Savia’ sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, reticulate, vasicentric, unilateral, or banded, or absent. Tyloses sometimes abundant. Sieve tube plastids S type?; sieve tubes with non-dispersive protein bodies (Bischofia). Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, unilacunar with one trace). Laticifers absent (resins excreted in Spondianthus and Uapaca). Silica bodies present or absent. Prismatic calciumoxalate crystals abundant; styloids and/or other types of crystals present or absent.

Trichomes Hairs unicellular or multicellular, usually simple (rarely stellate or lepidote); glands rarely present (on leaves).

Leaves Usually alternate (usually spiral, sometimes distichous; rarely opposite or whorled), usually simple (in Bischofia trifoliolate or sometimes pinnately compound), entire, sometimes coriaceous, with conduplicate or involute ptyxis; leaves on orthotropic branches spiral and often reduced (sometimes scale-like), on plagiotropic branches usually distichous, large and photosynthetic (phyllanthoid branching, branches resembling compound leaves). Stipules often scale-like, membranous, early caducous (rarely absent); leaf sheath absent. Petiole in Bischofia pulvinate. Petiole vascular bundle transection? Venation pinnate, brochidodromous or eucamptodromous. Stomata usually paracytic (sometimes anisocytic). Cuticular wax crystalloids? Abaxial side of lamina in Hymenocardia densely beset with red glandular dots. Epidermis sometimes with mucilaginous idioblasts. Domatia, as pockets or pits, present in some genera. Secretory cavities absent. Leaf margin usually entire (sometimes serrate, rarely with glands; in Bischofia with caducous teeth). Extrafloral nectaries often present on stipules, petiole and/or lamina.

Inflorescence Terminal or axillary, fasciculate, raceme-, spike- or catkin-like, cymose,or racemose inflorescence, or flowers solitary (usually female flowers). Inflorescenses (with very short branches) often arising in axils of leaves on plagiotropic shoots (phyllanthoid branching). Bracts sometimes involucrate (occasionally showy; in Uapaca pseudanthium).

Flowers Actinomorphic, small. Hypogyny. Sepals two to eight (to twelve), usually with imbricate or valvate (sometimes open) aestivation, free or connate at base (sometimes absent in female flowers). Petals (three to) five (or six), free, usually very small (sometimes absent). Nectariferous disc extrastaminal or interstaminal, annular or consisting of separate parts (sometimes central or absent), sometimes? with glands.

Androecium Stamens two or several (in Phyllanthus up to 15; in Tacarcuna 14 to 18; in Lingelsheimia c. 15 to c. 35). Filaments free or more or less connate, free from tepals. Anthers basifixed to dorsifixed, often versatile?, tetrasporangiate, usually extrorse (sometimes introrse), longicidal (dehiscing by usually longitudinal, sometimes transverse, slits; rarely apical, with pore-like slits). Tapetum secretory, with binucleate to quinquenucleate (Bischofia) cells. Female flowers often with staminodia; male flowers rarely with staminodia (fertile stamens of male flowers in Uapaca sometimes alternating with staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tri- to tetra- (rarely penta-)colporate (sometimes porate or inaperturate; sometimes with diploporate apertures; rarely pantosyncolpoidorate with up to c. 60 apertures; in Phyllanthus polypantoporate), shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, usually reticulate or smooth (rarely echinate or spinulate).

Gynoecium Pistil composed of one carpel (pseudomonomery) or two to five (to 15) connate carpels. Ovary superior, unilocular or bi- to quinquelocular (often? divided by pseudophragmata). Style single (sometimes lateral-subterminal), simple or branched, or stylodia two to four. Stigmas usually bilobate, fimbriate, with adaxial furrow, papillate?, Wet type. Male flowers often with shortly stipitate, peltate and entire or split pistillodium.

Ovules Placentation apical. Ovules (one or) two per carpel, anatropous, anacampylotropous or hemianatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument usually two to four (rarely five or more) cell layers thick. Inner integument usually two or three (rarely up to ten or more) cell layers thick. Megasporangium at least ten cell layers thick, protruding, persistent. Placental obturator present between stylar canal and micropyle. Hypostase present. Parietal tissue often protruding through micropyle. Nucellar beak present. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad or solanad.

Fruit Usually a septicidal capsule with persistent columella (disintegrating into six valves or three mericarps each with two valves), or a bi- to quadripartite schizocarp with samaroid mericarps (rarely a berry or a single- to six-seeded drupe).

Seeds Aril? Carunculus usually absent (sometimes present, more or less rudimentary). Seed coat usually exotegmic. Testa sometimes vascularized (in Aporosa etc. sarcotestal). Tegmen usually two to five (rarely up to 20 or more) cell layers thick. Exotegmen with (sometimes also radially elongate) ribbon-shaped cells with often sinuous cell walls, usually fibrous (sometimes palisade), often tracheoidal (in Didymocistus and Hymenocardium collapsed tracheoidal, also large endotegmic cells with tannins), sometimes (in, e.g., Aporosa) with sclerotic cells (exotegmen absent in Poranthera). Perisperm not developed. Endosperm usually copious (sometimes sparse or absent), oily. Embryo straight or curved, well differentiated, with or without chlorophyll. Cotyledons two. Germination?

DNA Mitochondrial maturase gene matR lost in Croizatia and Lachnostylis. Mitochondrial coxI intron present in Breynia and Phyllanthus.

Phytochemistry Cyanidin, delphinidin, cucurbitacins and other triterpenes, non-hydrolyzable tannins (geraniin), tropane alkaloids (phyllalbin), pyrrolizidine alkaloids and other alkaloids (securinine, phyllantine, phyllochrisine, etc.), and tyrosine-derived cyanogenic compounds present. Ellagic acid not found. Flavonols? Saponins? Aluminium accumulated in some species.

Use Ornamental plants, timber, edible fruits, medicinal plants, fish poison.

Systematics Phyllanthaceae are sister to Picrodendraceae.

Phyllanthaceae are provisionally subdivided into Phyllanthoideae (the ’Fasciculate clade’) and Antidesmatoideae (the ’Tanniniferous clade’) by Kathriarachchi & al. (2005).

Phyllanthoideae Beilschm. in Flora 16(Beibl. 7): 61, 109. 14 Jun 1833 [‘Phyllantheae’]

33/>1.660. Andrachne (22; the Mediterranean, southwestern Asia, western North America, the West Indies), Meineckia (c 30; southwestern and northeastern tropical Africal, Madagascar, Socotra, southern Arabian Peninsula, southern India, Sri Lanka, Assam, tropical America), Notoleptopus (1; N. decaisnei; Malesia to New Guinea and northern Australia), Poranthera (14; Australia, Tasmania, New Zealand), Pseudophyllanthus (1; P. ovalis; southern Africa), Phyllanthopsis (2; P. arida, P. phyllanthoides; Texas, Mexico), Actephila (c 35; southern China, tropical Asia to eastern Queensland, eastern New South Wales and islands in the Pacific), Leptopus (10; the Caucasus, northern Iran and the Himalayas to China and Southeast Asia), Heywoodia (1; H. lucens; tropical East Africa, KwaZulu-Natal, Eastern Cape), Astrocasia (6; A. austinii, A. diegoae, A. jacobinensis, A. neurocarpa, A. peltata, A. tremula; southern Mexico, Central America, the West Indies, Colombia, Venezuela), Chascotheca (1; C. neopeltandra; Cuba, Hispaniola), Dicoelia (1; D. beccariana; West Malesia), Wielandia (13; southeastern Kenya, Madagascar, the Comoros, the Seychelles), Margaritaria (14; tropical regions on both hemispheres), Lingelsheimia (6; L. abbayesii, L. ambigua, L. fiherenensis, L. frutescens, L. manongarivensis, L. sylvestris; Central Africa to Tanzania, Madagascar), Flueggea (16; western Mediterranean, Turkey, tropical and subtropical regions on both hemispheres), Plagiocladus (1; P. diandrus; western Central Africa), Richeriella (1; R. gracilis; northeastern India, Hainan, Thailand, West and Central Malesia; in Flueggea?), Heterosavia (4; H. bahamensis, H. erythroxyloides, H. laurifolia, H. maculata; Florida Keys, Central America, Cuba, Bahamas, Grand Cayman), Phyllanthus (>1.250; tropical and subtropical regions on both hemispheres), Securinega (5; S. antsingyensis, S. capuronii, S. durissima, S. perrieri, S. seyrigii; Madagascar, the Mascarene Islands), Lachnostylis (3; L. bilocularis, L. hanekomii, L. hirta; Western and Eastern Cape), Amanoa (16; tropical Africa, Madagascar, tropical America), Keayodendron (1; K. bridelioides; Ivory Coast to Cameroon), Savia (2; S. dictyocarpa, S. sessiliflora; southern Mexico, the West Indies, Venezuela, Paraguay, southern Brazil), Gonatogyne (1; G. brasiliensis; São Paulo in Brazil), Croizatia (5; C. brevipetiolata, C. cimalonia, C. naiguatensis, C. neotropica, C. panamensis; Panamá, Venezuela), Discocarpus (4; D. essequeboensis, D. gentryi, D. pedicellatus, D. spruceanus; northeastern South America), Tacarcuna (3; T. amanoifolia, T. gentryi, T. tachirensis; Panamá, Colombia, Venezuela, Peru), ’Cleistanthus’ (c 140; tropical regions in the Old World; paraphyletic), Pseudolachnostylis (1; P. maprouneifolia; tropical and southern Africa), Pentabrachion (1; P. reticulatum; Cameroon, Gabon), Bridelia (c 50; tropical Africa and Madagascar to northern Australia and islands in the Pacific). – Tropical and subtropical regions on both hemispheres, few species in the Mediterranean, Turkey, the Caucasus, southwestern Asia and western North America. Pistil composed of two to six (to 15) connate carpels. – Croizatia has five petals, extrastaminal disc and bifid style.

Antidesmatoideae Hurus. in J. Fac. Sci. Univ. Tokyo, ser. 3, Bot., 6: 321, 340. 15 Aug 1954

20/435–440. Bischofia (2; B. polycarpa: central and southeastern China; B. javanica: from India to eastern Asia, Melanesia and Polynesia to Samoa and Niue), Spondianthus (1; S. preussii; tropical East and Central Africa), Uapaca (c 50; tropical Africa, Madagascar), Protomegabaria (3; P. macrophylla, P. meiocarpa, P. stapfiana; tropical West and Central Africa), Richeria (5; R. australis, R. dressleri, R. grandis, R. obovata, R. tomentosa; tropical South America), Aporosa (75–80; tropical Asia to Solomon Islands), Maesobotrya (c 20; tropical Africa), Baccaurea (c 45; tropical Asia, islands in western Pacific; incl. Distichirhops?, Nothobaccaurea?), Distichirhops (3; D. megale, D. minor, D. mitsemosik; Borneo, New Guinea; in Baccaurea?), Nothobaccaurea (2; N. pulvinata, N. stylaris; Solomon Islands, Fiji; in Baccaurea?), Celianella (1; C. montana; sandstone tepuís in southern Venezuela), Jablonskia (1; J. congesta; northern South America), Hieronyma (c 20; southern Mexico, Central America, the West Indies, tropical South America), Leptonema (2; L. glabrum, L. venosum; Madagascar), Apodiscus (1; A. chevalieri; tropical West Africa), Martretia (1; M. quadricornis; tropical West and Central Africa), Antidesma (c 170; tropical and subtropical regions in the Old World), Thecacoris (c 25; tropical Africa, Madagascar), Hymenocardia (6; H. acida, H. heudelotii, H. lyrata, H. punctata, H. ripicola, H. ulmoides; tropical and southern Africa, Southeast Asia, Sumatra), Didymocistus (1; D. chrysadenius; tropical South America). – Tropical and subtropical. Petals often absent. Pistil composed of two to five connate carpels, or one carpel. Tannins present.

Unplaced Phyllanthaceae

Ashtonia (2; A. excelsa, A. praeterita; the Malay Peninsula, Borneo), Chonocentrum (1; C. cyathophorum; Amazonia)?

Cladogram of Phyllanthaceae based on DNA sequence data mainly according to Wurdack & al. (2004). The topology of the Poranthereae clade follows Vorontsova & al. (2007) and Vorontsova & Hoffmann (2008).

Cladogram of Phyllanthaceae based on DNA sequence data (Kathriarachchi & al. 2005).

Micrantheaceae J. Agardh, Theoria Syst. Plant.: 182. Apr-Sep 1858 [‘Micrantheae’]; Pseudanthaceae Endl. in H. Pfeiffer, Nomencl. Bot. 2(2): 852. 3 Oct 1873; Androstachyaceae Airy Shaw in Kew Bull. 18: 250. 8 Dec 1965 [‘Androstachydaceae’]; Paivaeusaceae (U. Köhler et G. L. Webster ex G. L. Webster) A. Meeuse, Euphorbiaceae: 30. 1990

Distribution Tropical and southern Africa, Madagascar, southern India, Sri Lanka, West Malesia, New Guinea, Australia, Tasmania, New Caledonia, southwestern United States to Argentina, with their highest diversity in Australia.

Fossils Rosenkrantzia picrodendroides from the Danian (Early Paleocene) of western Greenland, comprises fruits that resemble those in modern Picrodendraceae.

Habit Usually monoecious or dioecious (rarely bisexual), evergreen or deciduous trees or shrubs, perennial herbs; many species are stem succulents or twining.

Vegetative anatomy Phellogen ab initio superficial? Vessel elements with simple or scalariform perforation plates; lateral pits alternate or opposite, simple or bordered pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, winged-aliform, confluent, vasicentric, unilateral, or banded, or absent. Tyloses often frequent. Sieve tube plastids S type? Nodes? Laticifers absent. Heartwood with gum-like substances. Silica bodies sometimes abundant. Calciumoxalate as acicular crystals, styloids, crystal sand and other crystal types present in some species. Wood parenchyma and/or wood ray cells with prismatic crystals.

Trichomes Hairs unicellular or multicellular, uniseriate, simple; glandular hairs?

Leaves Alternate (usually spiral, sometimes distichous), opposite or verticillate, simple or palmately compound, entire or lobed, or absent, with ? ptyxis. Stipules petiolar or cauline (sometimes small, usually intrapetiolar; in Androstachys ochreate, enclosing terminal bud) or absent; leaf sheath absent. Colleters often present in leaf axils. Petiole vascular bundle transection? Venation palmate or pinnate. Stomata paracytic; subsidiary cells present on top of guard cells. Cuticular wax crystalloids? Epidermis usually with mucilaginous idioblasts. Secretory cavities absent. Leaf margin serrate (teeth with caducous apex) or entire. Foliar glands usually absent (sometimes with glands along leaf margins).

Inflorescence Terminal or axillary, catkin-like thyrse or fasciculate inflorescence (males), or flowers solitary axillary (females).

Flowers Actinomorphic, small. Hypogyny. Sepals in male flowers (two to) four to eight (to 13), with imbricate aestivation, spiral, free, or absent; sepals in female flowers (three or) four to eight (to 13), with valvate or imbricate aestivation, whorled, in some genera unequally sized, free, or absent. Petals absent. Nectariferous disc central or interstaminal, or absent (stamens rarely inserted in cavities on disc).

Androecium Stamens two to more than 100, alternate (sometimes on prolonged receptacle). Filaments usually free (rarely connate at base), free from tepals. Anthers dorsifixed, versatile?, apiculate, tetrasporangiate, extrorse to latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 4–12-colporate, 5–6-porate to zonaperturate or hexacolporoidate or inaperturate (with up to c. 60 very narrow apertures), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, spinulate, echinate or verruculate.

Gynoecium Pistil composed of two to four (or five) connate carpels. Ovary superior, usually bilocular to quadrilocular (or quinquelocular). Style usually single, simple, or stylodia two to four (or five), free. Stigmas stout, sometimes lobate, papillate?, usually Dry (sometimes Wet) type. Male flowers often with pistillodium.

Ovules Placentation axile to apical. Ovules usually two (in Scagea one) per carpel, anatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle usually bistomal (in Austrobuxus endostomal). Outer integument five or six cell layers thick. Inner integument three to six cell layers thick. Funicular obturator present between stylar canal and micropyle. Hypostase present. Nucellar cap present. Nucellar beak often present. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad?

Fruit Usually a septicidal and/or loculicidal capsule with persistent column, or a tripartite schizocarp (rarely a berry or a single- or two-seeded drupe with thin fleshy orange pericarp containing bitter juice).

Seeds Aril present or absent. Carunculus usually present (absent in, i.a., Podocalyx). Testa? Exotegmen cuboid or fibrous (exotegmen in Oldfieldia palisade, subprocumbent). Mesotegmen in Oldfieldia thickened. Endotegmen usually absent (in Oldfieldia two-layered, with band-shaped cell wall thickenings). Perisperm not developed. Endosperm usually copious, oily (sometimes absent; in Picrodendron ruminate). Embryo curved, well differentiated, with chlorophyll. Cotyledons two, sometimes strongly plicate (in Picrodendron lobate). Germination?

Phytochemistry Insufficiently known. Flavones, oleanolic acids, triterpenes and toxic picrotoxans (e.g. hyaenanchin) present. Saponins and cyanogenic compounds not found.

Kunth in von Humboldt, Bonpland et Kunth, Nov. Gen. Sp. 1: ed. 4°: 246. Mai 1816 [’Podostemeae’], nom. cons.

Marathraceae Dumort., Anal. Fam. Plant.: 60, 62. 1829 [‘Marathrineae’]; Marathrales Dumort., Anal. Fam. Plant.: 60. 1829 [‘Marathrarieae’]; Podostemales Lindl., Nix. Plant.: 17. 17 Sep 1833; Philocrenaceae Bongard, Mém. Acad. Imp. Sci. St. Pétersb., sér. 6, Math. Nat. 3(2): 87. 13 Jun-13 Jul 1834; Podostemineae Engl., Syllabus, ed. 2: 124. Mai 1898; Tristichaceae J. C. Willis in Bot. J. Linn. Soc. 43: 51. 15 Mai 1915; Podostemopsida G. Cusset & C. Cusset in Bull. Mus. Natl. Nat. Paris, Sect. B, Adansonia, sér. 4, 10: 210. 14 Oct 1988; Podostemanae R. Dahlgren ex Reveal in Novon 2: 236. 13 Oct 1992 [‘Podostemonanae’]

Distribution Tropical, subtropical and warm-temperate regions on both hemispheres.

Fossils Uncertain. Fossil leaves under name of Nitophyllites zaisanica have been found in Late Eocene layers in Russia and interpreted as Podostemaceae.

Habit Bisexual, usually annual (sometimes perennial) herbs. Aquatic (rheophytes) in rapidly running fresh water. Stems reduced or prolonged, simple or branched, sympodial, sometimes dimorphic or developed during anthesis only, often with specialized thalloid adventitious roots on the lower side and attached to the substrate by adhesive usually unicellular hooked hairs, hapters, excreting viscid polysaccharides (attaching to biofilm produced by cyanobacteria in the substrate). Adventitious roots dorsiventrally or laterally flattened, filiform, strap-shaped or discoid, creeping or partially floating, sometimes ephemeral or absent. Branching extra-axillary. Leaves inserted on prolonged stems or on procumbent, often discoid, stems. Podostemaceae with ribbon-like roots bear opposite branches, whereas species with foliose or crustose species develop single endogenous shoots from their upper surface (sometimes from cortex).

Vegetative anatomy Main root early withering. Adventitious roots photosynthesizing, usually endogenous (sometimes exogenous). Shoots nearly always arising as endogenous lateral buds from roots (in Cladopus both exogenous and endogenous lateral buds and lateral adventitious roots). Phellogen absent. Secondary lateral growth absent. Typical xylem and phloem elements (vessels, tracheids, libriform fibres, sieve tubes etc.) usually absent. Wood rays absent. Parenchyma? Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Laticifers and/or resinous cells present in many Podostemoideae. Air canals and aerenchyma rare. Silica bodies frequent in epidermis (and in subepidermal layers) in many species. Calciumoxalate crystals?

Leaves Alternate (usually distichous or tristichous, sometimes spiral, rarely tetrastichous to hexastichous), opposite or absent, scale-like to hair-like, simple or compound, entire or lobed, with ? ptyxis. Stipules petiolar or absent; leaf sheath simple or double, with lobes sometimes prolonged into stipule-like appendages. Petiole vascular bundle transection? Venation pinnate. Stomata at least usually absent. Cuticular waxes absent. Schizogenous secretory canals present. Leaf margin entire.

Inflorescence Terminal or axillary, cymose or racemose of various shape, or flowers solitary. Floral buds often endogenous. Usually only one floral bud naked (Weddellinoideae, some Tristichoideae) or surrounded by vascularized cupule (other Tristichoideae), or entirely enclosed within tubular or saccate membranous and non-vascularized spathella formed by two connate leaves or part of leaves (Podostemoideae).

Flowers Actinomorphic or zygomorphic (sometimes inverted in bud). Hypogyny. Tepals (four or) five (or six), with imbricate aestivation, sepaloid (Tristichoideae, Weddellinoideae) or two to c. 25, thin (staminodial?), usually alternating with stamens (Podostemoideae; in flowers with two basally connate stamens sometimes with additional tepal at apex of andropodium), spiral or in complete or incomplete whorl, often restricted to one side of flower, or reduced to annular margin, or absent; when few then more or less connate. Nectary absent. Disc absent.

Androecium Stamens one, few or c. 40 in one or two complete whorls, or in one incomplete whorl, or restricted to one side of flower and constisting of one to three free stamens, or Y-shaped structure composed of andropodium with two (to seven) stamens (cf. Podostemum, referring to Y-shaped androecium). Filaments free (in Tulasneantha connate into tube) or connate at base into fascicles (rarely paired), free from tepals. Anthers basifixed to subbasifixed, non-versatile?, tetrasporangiate (usually with microsporangia in one row?), usually introrse to latrorse (when two whorls then inner extrorse), longicidal (dehiscing by longitudinal slits); connective sometimes prolonged. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis usually simultaneous (sometimes successive). Pollen grains pantoporate with up to 16 pores (Tristichoideae), tricolporate (Weddellinoideae), tricolpate to pentacolpate (Podostemoideae) or rarely inaperturate, usually shed as monads or dyads (in Diamantina tetrads), bicellular at dispersal. Exine tectate, with granular infratectum, usually echinate (in Weddellinoideae rugulo-areolate).

Gynoecium Pistil composed of two to five connate carpels; antepetalous when carpels as many as tepals. Ovary superior, usually bilocular or trilocular (in some Podostemoideae unilocular). Gynophore usually present. Style single, usually simple (stylodia in Diamantina two), or absent. Stigmas one to three, of various shape, often linear, papillate?, type? Pistillodium absent.

Ovules Placentation usually axile (sometimes free central, when ovary unilocular). Ovules usually numerous (sometimes two or several) per carpel, anatropous, bitegmic, tenuinucellar. Micropyle exostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Apical part of megasporangium – part of megasporangium below megasporocyte and surrounding developing megagametophyte – significantly elongating and extending far beyond inner integument. Megagametophyte development monosporous or disporous, quadrinucleate or quinquenucleate, tricellular, quadricellular or quinquecellular, Apinagia type, Dacraea, Podostemon or Polypleurum subtypes. Meiosis II taking place in both dyad cells (no cell division); micropylar cell reduced to cap-like structure above megagametophyte. Chalazal megaspore nucleus dividing resulting in polar nuclei or one or two antipodals, or entirely or partially degenerating. Micropylar megaspore nucleus dividing vertically and chalazal nucleus horizontally (Dacraea and Polypleurum types, polar cell micropylar), or micropylar megaspore nucleus dividing horizontally and chalazal nucleus vertically (Podostemon type, polar cell chalazal). Polar nuclei often absent. Four micropylar nuclei forming megagametophyte, chalazal part developing into multinucleate nucellar plasmodium, produced by hyponucellus (part of megasporangium below megagametophyte). Antipodal cells usually absent (sometimes one reduced antipodal cell present). Fertilization simple (double fertilization not occurring). Endosperm not developing. Megasporangium plasmodial after fertilization. Endosperm haustorium micropylar, usually as well developed suspensor haustorium. Embryogenesis solanad.

Fruit Usually a septicidal capsule (in Farmeria metzgerioides a berry or drupe?).

Seeds Aril absent. Exotesta thick-walled, usually mucilaginous. Endotesta? Tegmen unspecialized. Exotegmic cell walls sometimes lignified. Endotegmic cell walls lignified. Suspensor present. Perisperm not developed. Endosperm present. Embryo straight (fusiform?), chlorophyll? Cotyledons usually two, large (sometimes one). Hypocotyl and plumule usually absent. Germination phanerocotylar. Radicula ephemeral or absent.

Phytochemistry Insufficiently known. Isoprenylated and other xanthones usually present (not found in Tristichoideae). Salts often accumulated.

Systematics Podostemaceae are sister to Hypericaceae.

Tristichoideae Warming in Kong. Danske Vidensk. Natur. Math. Afl. 6(11): 53. 1901 [’Tristicheae’]

6/14–18. Tristicha (1; T. trifaria: tropical and southern Africa, Madagascar, Mauritius, India, tropical America); Indotristicha (3; I. malayana, I. ramosissima, I. tirunelveliana; India), Dalzellia (6; D. angustissima, D. gracilis, D. kailarsenii, D. ranongensis, D. ubonensis, D. zeylanica; Sri Lanka to southern China and Southeast Asia), Indodalzellia (1; I. gracilis; southern India), Cussetia (2; C. carinata, C. diversifolia; Thailand, Cambodia, Laos; possibly extinct); Terniopsis (1–5; T. malayana; southern China, Thailand, the Malay Peninsula). – Tropical Africa, Madagascar, tropical Asia and eastwards to northern Australia, tropical America. Root apical meristem sometimes on both adaxial and abaxial side. Roots sometimes without root cap. Spathella absent. Hypogyny. Tepals three, usually free (sometimes connate). Stamens (one to) three (stamen in Tristicha one, adaxial). Pollen grains pantoporate. Pistil composed of three sometimes connate carpels. Integument developing simultaneously. Capsule with strong ribs. Hypocotyl absent. Xanthones possibly absent. – Tristichoideae are sister-group to [Weddellinoideae+Podostemoideae]. Some species of Dalzellia have a cupule at pedicel base, developed from leafy shoots, the outer and inner integuments are developing together, and the megasporangium is coenocytic. Tristicha has an adaxial stamen and the median carpel may be abaxial. Cussetia is probably closely allied to Tristicha and Terniopsis.

[Weddellinoideae+Podostemoideae]

Pistil composed of two connate carpels. Ovary with apical septum.

Weddellinoideae (C. Cusset et G. Cusset) Engl. in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 18a: 28. 3 Mai 1930

1/1. Weddellina (1; W. squamulosa; northern South America). – Plant scaly. Flowers terminal, solitary. Tepals (four or) five (or six), single-veined. Stamens five to c. 25. Pollen grains smooth. Stigmas spherical. Integuments developing simultaneously. Megagametophyte development very variable. Capsule without ribs. Tegmic cells thick-walled. Hypocotyl present.

Podostemoideae Wedd. in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 17: 39, 43. 16 Oct 1873 [‘Podostemoneae’]

50/278–285. Diamantina (1; D. lombardii; Brazil); ‘Mourera’ (7; M. alcicornis, M. aspera, M. elegans, M. fluviatilis, M. glazioviana, M. schwackeana, M. weddelliana; northern South America; paraphyletic), Monostylis (1; M. capillacea; Brazil), Castelnavia (11; Brazil), Rhyncholacis (c 25; northern tropical South America), Apinagia (≤50; tropical South America), Jenmaniella (7; J. ceratophylla, J. fimbriata, J. guianensis, J. isoetifolia, J. jenmanii, J. tridactylitifolia, J. varians; northeastern South America), Noveloa (2; N. coulteriana, N. longifolia; Mexico, Central America), Wettsteiniola (3; W. accorsii, W. apipensis, W. pinnata; southern Brazil, northern Argentina), Marathrum (≤25; southern Mexico, Central America, the West Indies, northwestern South America), Autana (1; A. andersonii; Venezuela), Ceratolacis (2; C. erythrolichen, C. pedunculatum; Brazil), Cipoia (2; C. inserta, C. ramosa; Brazil), Lonchostephus (1; L. elegans; Amazonian Brazil), Lophogyne (1; L. lacunosa; eastern central Brazil), Macarenia (1; M. clavigera; Colombia), Oserya (7; O. biceps, O. coulteriana, O. flabellifera, O. longifolia, O. minima, O. perpusilla, O. sphaerocarpa; southern Mexico, Central America, northern South America), Tulasneantha (1; T. monadelpha; western Brazil), Vanroyenella (1; V. plumosa; southwestern Mexico); Podostemum (17; eastern and southern United States, Mexico, Central America, the West Indies, tropical South America); Inversodicraea (4; I. congolana, I. garrettii, I. tenax, I. warmingiana; tropical Africa), Monandriella (1; M. linearifolia; Cameroon), Saxicolella (1; S. flabellata; tropical West and Central Africa), ’Ledermanniella’ (45–50; tropical and southern Africa; non-monophyletic), Letestuella (1; L. tisserantii; Kunene River in Namibia), Stonesia (2; S. fascicularis, S. gracilis; tropical West and Central Africa), Macropodiella (4–6; M. garrettii, M. hallaei, M. macrothyrsa, M. mildbraedii, M. pellucida, M. uoroensis; tropical West and Central Africa), Leiothylax (2; L. quangensis, L. warmingii; tropical Africa), Winklerella (1; W. dichotoma; tropical West Africa), Dicraeanthus (4; D. africanus, D. ramosus, D. taylorii, D. zehnderi; tropical West and Central Africa), Djinga (1; D. felicis; Cameroon), Endocaulos (1; E. mangorense; Madagascar), Thelethylax (2; T. isalensis, T. minutiflora; Madagascar), Angolaea (1; A. fluitans; Angola), Paleodicraeia (1; P. imbricata; Madagascar), Sphaerothylax (2; S. abyssinica, S. algiformis; tropical and southern Africa, Madagascar), Maferria (1; M. indica; southwestern India), Zehnderia (1; Z. microgyna; Cameroon); Cladopus (10; southern China, southern Japan, Southeast Asia, Malesia to New Guinea and northeastern Queensland), Paracladopus (2; P. chantaburiensis, P. chiangmaiensis; Thailand); Hydrodiscus (1; H. koyamae; Laos), Hydrobryum (5–10; H. floribundum, H. griffithii, H. japonicum, H. koribanum, H. puncticulatum; southern India, eastern Nepal, Assam, China, southern Japan), Hanseniella (2; H. heterophylla, H. smitinandii; Thailand), Thawatchaia (1; T. trilobata; Thailand); Willisia (1–2; W. arekaliana, W. selaginoides; Kerala in southern India), ‘Zeylanidium’ (5; Z. barberi, Z. maheshwarii, Z. olivaceum, Z. sessile, Z. subulatum; India, Sri Lanka; non-monophyletic), Griffithella (1; G. pierrei; Western Ghats in India), Farmeria (1; F. metzgerioides; Sri Lanka), Polypleurum (4; P. dichotomum, P. elongatum, P. stylosum, P. wallichii; India, Sri Lanka, northeastern Thailand), Diplobryum (4; D. koyamae, D. minutale, D. ramosum, D. vientianense; Laos, southern Vietnam). – Tropical and subtropical regions on both hemispheres, few species in warm-temperate regions. Shoots usually without apical meristem. Root apical meristems usually on lower side of thallus (sometimes on both upper and lower side). Roots sometimes exogenous, sometimes without root cap. Leaves ensiform, bifacial, often distichous, without normal epidermis, endogenously developed. Stipules sometimes also abaxial relative to leaf (leaves dithecal: one leaf sheath apically directed, additional leaf sheath basally directed). Stomata absent? Flowers or floral fascicles enclosed by non-vascularized spathella (possibly formed by fusion of two foliar structures). Tepals two to c. 25, with narrow lobes, sometimes replaced by stamens. Stamens one to three (to numerous). Microsporogenesis sometimes successive, with tetragonal tetrads. Pollen grains, 3–5-colpate, shed in monads or dyads, usually calymmate. Pistil composed of three (to seven) connate carpels. Ovary sometimes unilocular. Gynophore usually present. Style short, simple or branched, with long branches. Outer integument first developing. Megasporangium amoeboid-periplasmodial prior to fertilization. Megagametophyte sometimes disporous (Polypleurum and Podostemon types). Polar nucleus degenerating. Double fertilization not occurring. Capsule with ribs. Hypocotyl and radicula usually absent (hypocotyl present in Zeylanidium olivaceum). – Diamantina is sister to the remaining Podostemoideae (Kato & al. 2009; Ruhfel & al. 2011; Koi & al. 2012). In Old World Podostemoideae (“the Old World clades”) pollen grains are usually shed in dyads, whereas monads dominate in New World Podostemoideae (“the New World clade”).

Cladogram of Old World Podostemaceae based on DNA sequence data (Kato & al. 2009; Ruhfel & al. 2011).

Meisner, Plant. Vasc. Gen., Tab. Diagn. 345, Comm.: 258. 13-15 Feb 1842 [’Putranjiveae’]

Distribution Pantropical, southeastern Africa, Madagascar, subtropical East Asia, northern and eastern Australia, New Caledonia.

Fossils Uncertain. Fossil pollen grains attributed to Putranjivaceae have been reported from the Paleocene (Muller 1981).

Habit Usually dioecious (some species of Drypetes monoecious or polygamodioecious), evergreen trees or shrubs.

Vegetative anatomy Phellogen? Vessel elements with simple or scalariform perforation plates; lateral pits alternate, simple pits. Bordered intervessel pits abundant. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal reticulate, scalariform, or banded. Sieve tube plastids S type? Nodes? Latex and laticifers absent. Silica bodies present in parenchyma cells in some species. Wood parenchyma and/or wood ray cells with prismatic calciumoxalate crystals and silica bodies.

Leaves Alternate (distichous), simple, entire, often coriaceous, with ? ptyxis. Stipules present; leaf sheath absent. Petiole vascular bundle transection elliptic. Leaf base usually asymmetric. Venation pinnate; veins sometimes proceeding into transparent caducous teeth or spines. Stomata brachyparacytic with subsidiary cells overlying guard cells Cuticular wax crystalloids? Secretory cavities absent. Leaf margin serrate (sometimes spinose-serrate) or entire.

Inflorescence Axillary, fascicle (female inflorescences in Putranjiva one- to three-flowered).

Flowers Actinomorphic, small? Hypogyny. Sepals four or five (to seven), usually with imbricate (in female flowers of some species of Drypetes open) aestivation, usually free (in Putranjiva connate in lower part), in female flowers usually caducous. Petals absent. Nectary usually present (absent in Putranjiva). Disc annular (rarely absent), sometimes lobate (in Drypetes madagascariensis trilobate), with alternisepalous lobes, or cup-shaped, in male flowers central and encircling rudimentary gynoecium.

Androecium Stamens (two to) four to 20 (to more than 50 in Drypetes longifolia), diplostemonous in some species of Drypetes. Filaments usually free from each other and from tepals. Anthers usually basifixed (sometimes slightly dorsifixed), usually non-versatile, tetrasporangiate, usually extrorse or latrorse (rarely introrse), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate? infratectum, perforate or reticulate, usually smooth.

Gynoecium Pistil composed of two or three (to seven) connate carpels (sometimes seemingly one carpel; probably pseudomonomery). Ovary superior, synascidiate, unilocular to quadrilocular. Style usually single or absent (styles in Putranjiva two or three, connate at base), usually simple, expanded, usually short. Stigmas usually wide and flat (flap-shaped), simple or bifurcate (rarely multilobate, sometimes capitate), non-papillate (Drypetes) or papillate (Putranjiva), type? Pistillodium, very small, usually present (rarely absent) in male flowers.

Ovules Placentation axile. Ovules two per carpel, anatropous, pendulous, epitropous, usually bitegmic (in Drypetes macrostigma unitegmic with inner integument lost), weakly crassinucellar (with two parietal cell layers) or incompletely tenuinucellar. Micropyle endostomal or bistomal (sometimes exostomal; variation extensive). Outer integument three to seven cell layers thick, with few discrete vascular bundles. Inner integument five to nine (or more) cell layers thick, multiplicative. Single integument in Drypetes macrostigma six to nine cell layers thick. Obturator placental, stout, formed from funicular tissue near micropyle. Hypostase weakly differentiated in older ovules. Endothelium present. Parietal tissue approx. two cell layers thick. Archespore usually bicellular or tricellular (rarely quadricellular or quinquecellular). Megasporangium approx. two cell layers thick, early disintegrating. Nucellar beak absent. Nucellar cap absent. Megagametophyte monosporous, Polygonum type. Antipodal cells early degenerating. Endosperm development ab initio nuclear. Endosperm haustoria absent? Embryogenesis?

Fruit Usually a unilocular to quadrilocular drupe, often with persistent stigmas.

Seeds Aril absent. Carunculus absent. Testa vascularized. Sarcotesta thin or absent. Exomesotesta sclereidal. Endotesta? Tegmen multiplicative (six to more than 24 cell layers thick). Exotegmen sometimes fibrous. Exotegmic cells usually isodiametric, cuboidal, sometimes longitudinally elongate. Mesotegmen and endotegmen collapsing. Perisperm not developed. Endosperm copious, oily. Embryo straight, chlorophyll? Cotyledons two. Germination phanerocotylar?

Phytochemistry Insufficiently known. Biflavonoyls, cucurbitacins and other triterpenes, and glucosinolates (mustard oil glycosides) present. Flavonols? Ellagic acid? Saponins? Calciumoxalate present at least in floral organs.

Distribution Central America, the West Indies, tropical South America, with their largest diversity in Amazonia.

Habit Usually bisexual or androdioecious (sometimes polygamomonoecious, dioecious, or polygamodioecious), evergreen trees, shrubs or lianas.

Vegetative anatomy Mycorrhiza absent. Phellogen? Cortical vascular bundles absent. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids and fibres with bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, or vasicentric, or absent. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Cortex with cristarque cells. Parenchyma with secretory cells and druses. Heartwood in some species with resinous substance. Silica bodies or prismatic calciumoxalate crystals present in some species.

Leaves Opposite or verticillate, usually simple (in Froesia and Touroulia pinnately compound), entire or pinnately lobed, with ? ptyxis. Stipules usually interpetiolar, large, rigid or foliaceous (in Froesia deeply split), persistent; leaf sheath absent. Petiole vascular bundle transection annular, often complex. Venation (impari-)pinnate; secondary veins coarse, densely spaced; tertiary veins paxillate, parallel to plumose-reticulate, densely spaced; leaf veins surrounded by very thick-walled fibres. Stomata anisocytic or paracytic. Cuticular wax crystalloids? Lamina with lysigenous secretory mucilage cavities. Mesophyll with calciumoxalate as druses and single prismatic crystals. Leaf margin serrate or crenate.

Inflorescence Terminal or axillary, basically thyrsoid (determinate thyrse).

Flowers Usually actinomorphic (rarely somewhat zygomorphic). Hypogyny. Sepals four or five, with imbricate aestivation, unequal, free. Petals four or five (to eight), usually with imbricate (sometimes contorted) aestivation, free. Nectary absent. Disc absent.

Androecium Stamens twelve to more than 170. Filaments free or connate at base, free from or adnate at base to petals. Anthers dorsifixed, non-versatile, tetrasporangiate, introrse (thecae separate) or latrorse (thecae arranged side by side, in Froesia), longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, microperforate.

Gynoecium Pistil composed of two, three or seven to 14 (in Lacunaria four to 14) carpels usually paracarpously connate below (in Froesia three, free, apocarpy). Ovary superior, bilocular, trilocular or septa to 14-locular (in Lacunaria quadrilocular to 14-locular). Stylodia two, three or seven to 14, free. Stigmas obliquely peltate, type? Pistillodium often present in male flowers.

Ovules Placentation basal to axile. Ovules two per carpel, anatropous, ascending, bitegmic, tenuinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. ‘False endothelium’ present on surface of megasporangium. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A berry or berry-like single- to four-seeded valvate capsule (in Froesia follicles) with persistent calyx. Exocarp with lacunae.

Seeds Aril absent. Seeds usually densely tomentose (in Froesia glabrous or almost glabrous). Testa? Tegmen? Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, thick. Germination cryptocotylar.

Rafflesiales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 18. Sep-Oct 1835 [‘Rafflesiaceae’]; Rafflesianae Thorne ex Reveal in Phytologia 79: 71. 29 Apr 1996

Distribution Assam, Bhutan, southernmost China, Burma, Thailand, Indochina, West Malesia to the Philippines.

Habit Usually dioecious (in Rhizanthes lowii and R. zippelii also bisexual), achlorophyllous perennial herbaceous endophytic stem and root holoparasites without rhizome or ordinary roots, parasitizing species of Tetrastigma (Vitaceae). Flowers evil-smelling.

Vegetative anatomy Mycorrhiza absent. Vegetative tissue consisting of mostly uniseriate parenchymatous filaments probably capable of intrusive intercellular growth and transfer of nutrient and water (possibly derived from laticifers with similar type of growth and frequently present in Euphorbiaceae). Hypha-like cellular threads invading host plant and forming endophytic system inside roots and/or stem. Phellogen absent. Secondary lateral growth absent. Vessel elements probably absent. Imperforate tracheary xylem elements libriform fibres? (or absent?) Wood rays absent. Axial parenchyma absent? Sieve tube plastids S₀ type, without starch or protein inclusions. Nodes? Crystals?

Leaves Usually verticillate (sometimes opposite or alternate spiral), reduced and scale-like, or absent, with ? ptyxis. Stipules and leaf sheath absent. Venation? Stomata anomalous, with three or more guard cells (Rafflesia), or absent. Cuticular waxes absent. Leaf margin entire.

Inflorescence Flowers solitary. Floral shoots endogenous, bursting out through cortex of host plant.

Flowers Actinomorphic, medium-sized to very large (Rafflesia arnoldii with largest flower of all angiosperms, reaching almost 1 m across). Flower in Rafflesia and Rhizanthes surrounded by three whorls of scale-like bracts with five bracts in each. Epigyny. Tepals (sepals?) four to ten (to 16), usually with imbricate (in Rhizanthes valvate) aestivation, in one whorl (uniseriate; in Sapria biseriate?), with incurved margins, usually connate in lower part, in Rafflesia and Sapria inserted around thin annular horizontal central tissue, diaphragm (homologous to corona or corolla?), on apex of perianth tube; apex of central column in Rafflesia enlarged and disc-shaped; apices of 16 tepals in Rhizanthes inserted inside cavity in central column (diaphragm absent). Base of perianth tube in Rafflesia covered by special outgrowths, ramenta, on adaxial side; diaphragm in Sapria covered by ramenta. Nectary present at stylar base or absent. Androgynophore present?

Androecium Stamens twelve to more than 50, inserted on and adnate to ring immediately below (disc-shaped) apex of central column. Filaments absent. Anthers basifixed, non-versatile, disporangiate (monothecal) to polysporangiate, connate into annular synandrium, extrorse, poricidal (dehiscing by apical pore). Tapetum secretory? Female flowers with staminodia (rudimentary anthers).

Pollen grains Microsporogenesis successive. Pollen grains inaperturate, shed as monads, bicellular at dispersal. Exine intectate; pollen surface smooth?

Gynoecium Pistil composed of three to ten connate carpels; carpellary margins occluded through postgenital fusion and secretion. Ovary inferior, unilocular. Style single, simple, very short. Stigma in Rafflesia and Rhizanthes as annular structure on outer margin or lower side of apex of (disc-shaped) central column, papillate, type? Male flowers in Rhizanthes with pistillodium (rudimentary ovary).

Ovules Placentation laminar-parietal. Ovules numerous per ovary, anatropous, unitegmic or bitegmic, tenuinucellar. Micropyle usually endostomal (rarely exostomal). Outer integument one cell layer thick or absent. Inner integument ? cell layers thick. When unitegmic, then swelling present on chalaza (representing second integument?). Nucellar cap present. Megasporangial epidermis persistent. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad or solanad?

Seeds Aril absent. Seed consisting of two different parts; part covered by testa not enclosing embryo. Exotesta as basal chalazal appendage. Endotesta hard. Exotegmic cell walls with U-shaped thickenings. Endotegmic cell walls with thickenings. Perisperm not developed. Endosperm single-layered, sparse. Embryo rudimentary, undifferentiated (with few cells in few tiers), without chlorophyll? Cotyledons two. Germination phanerocotylar?

Persoon, Syn. Plant. 2: 2. Nov 1806 [’Rhizophorea’, ’Rhizophoreae’], nom. cons.

Rhizophorales Pers. ex Bercht. et J. Presl, Přir. Rostlin: 257. Jan-Apr 1820 [‘Rhizophoreae’]; Mangiaceae Raf., Fl. Tellur. 3: 73. Jan-Mar 1837 [‘Mangidia’]; Legnotidaceae (Endl.) Blume, Mus. Bot. Lugd.-Bat.: 126. Oct 1849 [‘Legnotideae’], nom. illeg.; Cassipoureaceae J. Agardh, Theoria Syst. Plant.: 246. Apr-Sep 1858 [‘Cassipoureae’]; Macarisiaceae J. Agardh, Theoria Syst. Plant.: 295. Apr-Sep 1858 [‘Macharisieae’]; Rhizophoranae (Pers.) Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Distribution Atlantic and Indian Ocean coasts of Africa, islands in the Indian Ocean and the Pacific, Atlantic and Pacific coasts of Central and South America, coasts in the West Indies, with their highest diversity in Madagascar and tropical Asia.

Fossils Pollen grains (resembling those in Bruguiera, Kandelia and Rhizophora) have been found in Eocene layers in France. Fossil Rhizophoraceae pollen is also known from Caribbean layers of Late Eocene age, and have been recorded from Oligocene and younger strata in Mexico, Venezuela, the Caribbean, Nigeria and Malesia (extant distribution in this area is possibly less than 11 My old). Eocene macrofossils (Ceriops cantiensis, Palaeobruguiera elongata, etc.) are known from Europe.

Habits Usually bisexual (rarely polygamomonoecious), evergreen trees or shrubs. Many species are mangrove trees. Pneumatophores (in Gynotrocheae and Rhizophoreae) and stilt roots often present.

Vegetative anatomy Root hairs absent in Rhizophoreae. Phellogen ab initio at least sometimes superficial. Vessel elements with simple and/or scalariform perforation plates; lateral pits scalariform, opposite or alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, lozenge-aliform, confluent, reticulate, scalariform, vasicentric, unilateral, or banded. Tyloses often frequent. Sieve tube plastids Pc (PV) type, with 20 or more square or polygonal protein crystalloids. Nodes ≥3:≥3, trilacunar or multilacunar with three or more leaf traces, often with split lateral vascular bundles. Heartwood often with ethereal substances. Prismatic calciumoxalate crystals frequent; acicular crystals, styloids, crystal sand and other types of crystals (in Paradrypetes raphides) often present.

Trichomes Hairs unicellular or sometimes multicellular, uniseriate, simple or stellate; glandular hairs often present.

Leaves Usually opposite (rarely verticillate), simple, entire, coriaceous, usually with involute (in Rhizophoreae supervolute) ptyxis. Stipules interpetiolar, large, sheathing, caducous, with basal adaxial gum-excreting colleters; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata paracytic, cyclocytic (Rhizophoreae) or anomocytic (rarely anisocytic). Cuticular wax crystalloids? Epidermis with or without mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts (with H-shaped and other sclereids). Leaf margin serrate, crenate or entire (in Paradrypetes spinose-serrate); leaf teeth theoid.

Inflorescence Usually axillary, fasciculate, raceme- or spike-like (flowers in some species of Bruguiera solitary axillary; inflorescence in Paradrypetes epipetiolar).

Flowers Actinomorphic. Pedicel articulated. Hypanthium-like structure usually present. Hypogyny (Macarisieae), epigyny or half epigyny. Sepals (three or) four or five (to 16), with valvate aestivation, usually carnose or coriaceous, persistent, free. Petals (three or) four or five (to 16), with contorted or inflexed aestivation, small, alternisepalous, usually hairy and carnose, often bilobate, usually with apiculate arista and fimbriate or with filiform appendages, often clawed, free. Subepidermal laticifers present. Nectariferous disc intrastaminal, inserted on ovary or “hypanthium”, or absent. Sepals and ovary with layer of hypodermal laticiferous cells (in Gynotroches and Pellacalyx as numerous secretory idioblasts).

Androecium Stamens eight to numerous, usually at least twice as many as petals (sometimes as many as petals), diplostemonous or obdiplostemonous. Filaments usually free (rarely connate at base), antepetalous, single or in fascicles of two to five stamens, free from tepals, usually inserted on abaxial side of nectariferous disc; each stamen or staminal fascicle usually enclosed by petal. Anthers dorsifixed, versatile, tetrasporangiate (in Rhizophora multilocellate due to transverse septa), introrse or latrorse, longicidal (dehiscing by longitudinal valve). Tapetum secretory. Staminodia absent. Rhizophoreae with secondary pollen display.

Pollen grains Microsporogenesis simultaneous or successive (Gynotrocheae). Pollen grains 3(–4)-colporate to 3(–4)-colporoidate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate or granular-columellate infratectum, punctate-rugulate, punctate, reticulate or smooth (in Paradrypetes echinate).

Gynoecium Pistil composed of (two or) three to five (to 20) connate antesepalous carpels; when two carpels, then lateral (in Rhizophora transverse). Ovary superior, inferior or semi-inferior, unilocular or multilocular, synascidiate. Style single, usually simple (in Gynotroches branched; absent in Paradrypetes). Stigma punctate, capitate or lobate, usually papillate, type? Pistillodium absent.

Ovules Placentation apical to axile. Ovules two to numerous per carpel, anatropous to hemianatropous (in Bruguiera and Carallia campylotropous), pendulous, epitropous?, bitegmic, usually crassinucellar (in Gynotroches and Pellacalyx tenuinucellar). Micropyle usually bistomal, Z-shaped (zig-zag) (sometimes endostomal). Outer integument usually three to six (in Carallia approx. ten) cell layers thick, in Rhizophoreae vascularized. Inner integument four to eight (to 20) cell layers thick. Obturator present (Macarisieae) or absent (Gynotrocheae, Rhizophoreae). Endothelium usually present. Parietal tissue one to three cell layers thick. Megasporocytes often several. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral. Endosperm development ab initio nuclear. Endosperm haustorium micropylar and chalazal (Ceriops). Embryogenesis?

Fruit A single- or many-seeded berry (Gynotrocheae), a drupe with one seed per locule, a capsule (Macarisieae, Crossostylis), or with hard pericarp and indehiscent (Rhizophoreae; seedling in Bruguiera dispersed together with fruit).

Seeds Seeds sometimes winged. Aril/carunculus exostomal or absent. Seed coat usually exotegmic. Testa sometimes multiplicative, indistinct (Rhizophoreae), or (in Macarisieae and Gynotrocheae) exotestal cells more or less enlarged, thick-walled. Endotesta sometimes crystalliferous. Tegmen usually thickened (absent in ripe seeds ofRhizophoreae), with invaginations. Exotegmen in Macarisieae and Crossostylis fibrous. Endotegmen crushed (Macarisieae) Perisperm not developed. Endosperm sparse or copious, oily. Embryo long or short, straight, well differentiated, with chlorophyll. Cotyledons usually two (rarely three or four), with unilacunar node. Germination phanerocotylar or viviparous. In mangrove trees (Rhizophoreae) vivipary, with much enlarged hypocotyl.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, afzelechin, myricetin, ethereal oils (in wood), ellagic acid, methylated ellagic acids, non-hydrolyzable tannins, proanthocyanidins (procyanidin, prodelphinidins), necin and tropane (hygrolin) alkaloids (oxytropanes, brugine, tropin, etc.), and pyrrolizidine alkaloids as 1-aminopyrrolizidine derivatives present. Saponins and cyanogenic compounds not found.

Systematics Rhizophoraceae are sister to Erythroxylaceae.

Macarisieae Baill., Hist. Plant. 6: 295, 302. Jan-Mai 1876

6/c 63. Paradrypetes (2; P. ilicifolia, P. subintegrifolia; western Amazonas, coastal Brazil); Sterigmapetalum (c 9; tropical South America), Cassipourea (c 40; tropical, eastern and southern Africa, Madagascar, the Mascarene Islands, southern India, Sri Lanka, tropical America), Macarisia (8; Madagascar), Anopyxis (3; A. ealaensis, A. klaineana, A. occidentalis; tropical Africa), Blepharistemma (1; B. serratum; southwestern India), Comiphyton (1; C. gabonense; Gabon to eastern Congo). – Tropical and southern Africa, Madagascar, southwestern India, Sri Lanka, tropical South America. Calcium oxalate crystals solitary. Stipules valvate. Leaf teeth often theoid. Hypanthium-like structure sometimes present. Sepals with open aestivation. Filaments often of two different lengths. Anthers latrorse; connectives often enlarged. Stigma capitate or punctate. Aril often present. Fruit in Cassipourea a capsule. Seeds sometimes winged at micropylar end. Exotegmen fibrous. – Paradrypetes may be sister to the remaining Macarisieae. Brazil. Dioecious trees. Raphides present. Lamina with long, Z-shaped intersecondary veins. Flowers small. Tepals in male flowers three or four. Nectary absent. Exine spinulate. Style absent. Placental obturator present. Fruit a drupe. Testa and tegmen vascularized. Endosperm copious, starchy. Cotyledons wide, plicate.

[Gynotrocheae+Rhizophoreae]

Stilt roots or pneumatophores present. Rootlets without root hairs. Leaves bijugate. Stipules imbricate. Hypanthium-like structure present. Epigyny. Obturator absent. Testa vascularized.

Gynotrocheae Engl., Syllabus, ed. 1: 147. Apr 1892

4/30–32. Carallia (11; Madagascar, tropical Asia to tropical Australia and Solomon Islands), Crossostylis (10–12; Melanesia, southwestern Polynesia), Gynotroches (1; G. axillaris; Burma, Southeast Asia, Malesia to New Guinea, Solomon Islands, the Caroline Islands and northern Queensland), Pellacalyx (c 8; tropical Asia). – Madagascar, tropical Asia to tropical Australia, islands in western Pacific. Stilt roots absent in Pellacalyx. Carpels often more numerous than sepals. Ovules sometimes up to eight per carpel, tenuinucellar. Outer integument two or three cell layers thick. Inner integument two to four cell layers thick. Archespore sometimes one. Fruit usually a berry (fruit in Crossostylis a capsule). Aril usually absent (present in Crossostylis). Exotesta mucilaginous, tanniniferous. Meso- and endotestal cells crystalliferous. Tegmen absent or fibrous to palisade. Exotegmen usually well developed (absent in Carallia); exotegmen in Crossostylis fibrous. Mesotegmen and endotegmen persistent. Cotyledons in Carallia and Pellacalyx involute.

Rhizophoreae Bartl., Ord. Nat. Plant.: 320. Sep 1830 [‘Rhizophorea genuina’]

4/19. Bruguiera (6; B. cylindrica, B. exaristata, B. gymnorhiza, B. hainesii, B. parviflora, B. sexangula; coasts of tropical East Africa and east to Samoa), Rhizophora (6; R. apiculata, R. mangle, R. mucronata, R. racemosa, R. samoensis, R. stylosa; tropical coasts on both hemispheres), Kandelia (2; K. candel, K. obovata; coasts of India and Bangladesh to Borneo, China and Kyushu in southern Japan), Ceriops (5; C. australis, C. decandra, C. pseudodecandra, C. tagal, C. zippeliana; coasts of East Africa and India to Queensland, Melanesia, Micronesia and northern to southeastern China). – Tropical coasts on both hemispheres. Stomata cyclocytic. Abaxial hypodermis present. Sclerenchymatous sheath of midvein absent or poorly developed. Leaf margin entire. Petals sometimes postgenitally fused above base. Anthers in Rhizophora locellate. Endothelium absent. Fruit single-seeded, indehiscent. Seed coat undifferentiated, vascularized. Tegmen not persistent. Endosperm usually overflowing and opening micropyle (not in Bruguiera). Embryo usually large. Seeds germinating on tree (viviparous). Cotyledons in Bruguiera and Rhizophora convolute; cotyledonary node trilacunar or multilacunar. Hypocotyl finally straightening.

Cladogram of Rhizophoraceae based on DNA sequence data and morphology (Schwarzbach & Ricklefs 2000). Paradrypetes is sister to Cassipourea (Wurdack & Davis 2009).

Mirbel, Elém. Physiol. Vég. Bot. 2: 905. 24-30 Jun 1815 [’Salicineae’], nom. cons.

Prockiaceae Bertuch, Taf. Allg. Naturgesch. Gewächs-Reich: Enum. 5, Syn. Tab. 3. 1801 [’Prockiae’]; Samydaceae Vent. in Mém. Cl. Sci. Math. Inst. Natl. France 1807(2): 149. 1808 [’Samydeae’], nom. cons.; Homaliaceae R. Br. in J. H. Tuckey, Narr. Exped. Zaire: 438. 5 Mar 1818 [’Homalinae’]; Samydales Vent. ex Bercht. et J. Presl, Přir. Rostlin: 227. Jan-Apr 1820 [‘Samydeae’]; Flacourtiaceae Rich. ex DC., Prodr. 1: 255. med Jan 1824 [‘Flacourtianeae’], nom. cons.; Blakwelliaceae T. Lestib., Botanogr. Élem.: 519. Jun 1826 [‘Blakwelliées’], nom. illeg.; Salicopsida Bartl., Ord. Nat. Plant: 92, 118. Sep 1830 [’Salicinae’]; Salicales Lindl., Nix. Plant.: 17. 17 Sep 1833 [‘Salicinales’]; Flacourtiales Rich. in C. F. P. von Martius, Consp. Regn. Veg.: 58. Sep-Oct 1835 [‘Flacourtianeae’]; Homaliales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 63. Sep-Oct 1835 [‘Homalineae’]; Flacourtiineae Engl., Syllabus, ed. 2: 154. Mai 1898; Scyphostegiaceae Hutch., Fam. Fl. Pl. 1: 229. 15 Jan 1926, nom. cons.; Scyphostegiales Croizat, Kirkia 15: 137. Sep 1994; Bembiciaceae R. C. Keating et Takht. in Bot. Žurn. 81(2): 85. Mai-Jun 1996; Poliothyrsidaceae (G. S. Fan) Doweld, Tent. Syst. Plant. Vasc.: xxxi. 23 Dec 2001; Caseariaceae Doweld, New Syllabus Pl. Fam.: 693. 2008

Distribution Mainly tropical and subtropical regions, some genera in temperate regions, few in Australia, absent from New Zealand; Salix and Populus mainly in temperate regions and some species of Salix only in arctic-alpine regions on the Northern Hemisphere.

Fossils Pseudosalix from the Eocene of Utah and Colorado is morphologically intermediate between Salix and genera formerly included in ‘Flacourtiaceae’ (e.g. Bennettiodendron, Idesia, and Polyothyris). Fossil pollen grains assigned to Casearia have been described from Oligocene layers.

Habit Usually bisexual (sometimes dioecious, e.g., Salix, Populus, Chosenia, and Scyphostegia), evergreen or deciduous trees or shrubs (some genera climbing; few species of Salix dwarf shrubs). Casearia sometimes with phyllanthoid branching: orthotropic branches with reduced spiral leaves, and plagiotropic branches sylleptic with normal leaves distichous.

Vegetative anatomy Ectomycorrhiza present in at least Salix, Populus and Chosenia. Phellogen ab initio superfical. Primary medullary rays narrow. Primary vascular tissue cylinder, without separate bundles. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, usually bordered (sometimes simple) pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma absent or very rare. Tyloses often abundant. Sieve tube plastids S type; sieve tubes with non-dispersive protein bodies? Nodes usually 3:3, trilacunar with three leaf traces (in Xylosma and some Casearia 1:1, unilacunar with one trace, in some species of Azara 2:2, bilacunar with two traces). Secondary phloem sometimes stratified into hard fibrous and soft parenchymatous layers (Salix, Populus, Chosenia?), in, e.g., Flacourtia and Xylosma with groups of large sclereids. Secretory cavities absent. Heartwood often with ethereal? substances. Prismatic calciumoxalate crystals abundant; acicular crystals, druses, styloids, crystal sand and other types of crystals often present.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple, furcate, stellate, peltate, lepidote, or vesicular, or absent; glandular hairs often present (transparent in, e.g., Abatia, Casearia, Ryania, and Zuelania).

Leaves Usually alternate (spiral or distichous; in Abatia opposite), simple, usually entire (sometimes lobed), sometimes coriaceous, with supervolute-curved or involute ptyxis. Stipules small to foliaceous, intrapetiolar, usually caducous (sometimes persistent; rarely absent); leaf sheath absent. Petiole vascular bundle transection arcuate or annular; petiole with wing bundles (petiole in Salix, Populus and Chosenia with unique arrangement of vascular strands as one or several closed cylinders of xylem and floem, in Populus often superimposed). Venation usually pinnate (rarely palmate), brochidodromous, campylodromous or actinodromous. Stomata anomocytic, paracytic or anisocytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type) or absent. Domatia as pits or hair tufts. Epidermis with or without mucilaginous idioblasts, with or without crystalliferous idioblasts. Cystoliths present (Homalium) or absent. Mesophyll with calciumoxalate as druses and single prismatic crystals, with or without sclerenchymatous idioblasts. Lamina sometimes gland-dotted. Leaf margin usually serrate (rarely entire, crenate or crispate), often with salicoid teeth (one vein running into tooth, tooth apex expanding into spheroid glandular structure). Extrafloral nectaries often present on petiole and/or lamina (usually pairwise on junction of petiole and lamina).

Inflorescence Terminal or axillary (in Bembicia epiphyllous and cone-shaped; in Phyllobotryon and Phylloclinium epiphyllous and solitary or few-flowered), usually cymose (sometimes racemose) of various shape (spike-, head- or catkin-like, fasciculate, panicle, spike, or catkin; flowers rarely solitary). Floral prophylls (bracteoles) present or absent. Telescoping floral bracts present in Scyphostegia.

Flowers Actinomorphic, often small. Hypanthium present in some genera. Usually hypogyny (rarely epigyny or half epigyny). Sepals three to eight (to c. 15), with imbricate or valvate aestivation, often persistent, usually free (sometimes connate at base; rarely petaloid; sometimes absent). Petals three to eight (to c. 15; in, e.g., some Scolopieae outnumbering sepals), with valvate or imbricate aestivation, usually free (rarely connate at base; sometimes absent); tepals rarely (in, i.a., Oncoba) spiral and hardly subdivided into calyx and corolla; tepals in Salix connate and modified into one or several nectaries (tepals sometimes absent). Corona in some genera present as scales, hairs or lobes on petal bases. Nectariferous disc extrastaminal or intrastaminal, as glands, scales or lobate annular disc (sometimes absent).

Androecium Stamens one to c. 40 (to more than 100), usually antesepalous (sometimes alternisepalous), in one or several whorls (sometimes in three to eight antepetalous fascicles). Filaments filiform, free or connate at least at base (in Scyphostegia entirely connate), free from tepals. Anthers dorsifixed or basifixed, non-versatile, tetrasporangiate, usually introrse (rarely extrorse), longicidal (dehiscing by longitudinal slits); connective sometimes apically prolonged. Tapetum secretory. Staminodia in Scyphostegia three, extrastaminal. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate or tricolpate to tricolporoidate (Salix; in Populus inaperturate), shed as monads, usually bicellular (rarely tricellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate or reticulate, verrucate, spinulate, or echinate.

Gynoecium Pistil composed of two to five (to 13) usually connate carpels (in Oncoba secondarily free); gynophore present in some species. Ovary usually superior (rarely inferior or semi-inferior), usually unilocular (in Scyphostegia unilocular in lower part, multilocular in upper part; rarely entirely multilocular). Style single, simple, persistent, or stylodia several, free or connate in lower parts (sometimes absent). Stigmas one or several, lobate, capitate or attenuate, commissural or carinal, non-papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation usually parietal or free central (sometimes intrusively parietal; rarely axile or basal; in Scyphostegia basal). Ovules usually two to more than 100 (rarely one) per ovary, usually orthotropous (sometimes anatropous or hemianatropous), ascending, usually bitegmic (sometimes unitegmic, with inner integument rudimentary), crassinucellar. Micropyle usually bistomal (rarely exostomal or endostomal), sometimes Z-shaped (zig-zag). Outer integument two to five cell layers thick. Inner integument two to five cell layers thick. Hypostase usually absent (present in Casearia). Parietal tissue five to seven cell layers thick, finally resorbed. Nucellar cap usually present, finally resorbed. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type), often protruding through micropyle. Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad or asterad.

Seeds Seeds often with aril (in Salix, Populus and Chosenia with arillar hair tuft at base: aril modified into hair tuft). Testa multiplicative, carnose (otherwise indistinct), or exotesta or endotesta (Oncoba) palisade. Exotegmen usually fibrous, lignified. Exotegmic cells is Dovyalis ribbon type. Endotegmen persistent? Perisperm usually not developed (in Scyphostegia thin). Endosperm usually copious (sometimes sparse, absent in Salix), oily. Embryo large, usually straight, well differentiated, with or without chlorophyll. Cotyledons two, plano-convex or flat. Germination phanerocotylar or cryptocotylar.

DNA Gene duplication (“the salicoid duplication”) present in Populus and Salix. Plastid gene rps16 lost in Populus (P. deltoides) and Salix (S. amygdaloides).

Phytochemistry Flavonols (kaempferol, quercetin), biflavonoids, biflavanoids, cyanidin, ellagic acid, tannins with proanthocyanidins and catechin, proanthocyanidins (prodelphinidins), phenolic glycosides (salicin, populin, etc.), gynocardins, saponins, acetophenones, arbutin, myo-inisitol, nigracin, and co-carcinogenic substances present. Cyclopentenoid cyanogenic glycosides not found. Aluminium accumulated in some species.

Use Ornamental plants, matches (Populus), baskets (withies from Salix), bioenergy (Salix), timber, protective plantations (against wind and erosion), fruits (Dovyalis), medicinal plants.

Systematics Salicaceae are probably sister to Lacistemataceae.

Salicaceae are under construction and the subdivision below is highly provisional.

Samydeae Vent. in B. C. J. Dumortier, Anal. Fam. Plant.: 18. 1829

14/240–245. Casearia (c 180; tropical regions on both hemispheres), Euceraea (3; E. nitida, E. rheophytica, E. sleumeriana; Venezuela, Guyana, Suriname, Brazil), Hecatostemon (1; H. completus; Venezuela, Guyana, Brazil), Laetia (9; Central America, tropical South America), Lunania (15; southern Mexico, Central America, Cuba, Jamaica, northern South America), Neoptychocarpus (3; N. apodanthus, N. chocoensis, N. killipii; tropical South America), Ophiobotrys (1; O. zenkeri; tropical West and Central Africa), Osmelia (4; O. gardneri, O. grandistipulata, O. maingayi, O. philippina; Sri Lanka, Malesia), Pseudosmelia (1; P. moluccana; the Moluccas), Ryania (9; tropical America), Samyda (11; Central America, the West Indies), Tetrathylacium (2; T. johansenii, T. macrophyllum; tropical America), Zuelania (1; Z. guidonia; southern Mexico, Central America, Venezuela); Trichostephanus (2; T. acuminatus, T. gabonensis; tropical West and Central Africa). – Tropical regions on both hemispheres. Bisexual (in Trichostephanus monoecious) trees or shrubs. Stipules usually caducous. Lamina often with pellucid glandular dots and lines. Leaf margin serrate or entire; leaf teeth theoid (e.g. Casearia) or salicoid. Hypanthium present. Sepals three to seven, connate at base (absent in Trichostephanus). Petals absent. Nectariferous disc present at adaxial side of sepal bases (calyx tube), lobate. Stamens three to twelve (sometimes more). Ovary superior to semi-inferior, unilocular. Style simple or distally trifid. Stigma trilobate or stigmas capitate. Ovules several per carpel. Fruit a carnose or coriaceous capsule (often berry-like), usually with persistent sepals, disc, filaments and style. Aril vascularized, often fimbriate. Exotegmic cells laterally flattened, crystalliferous. – Casearia is sometimes identified as sister-group with low support to the remaining Salicaceae. The position of Trichostephanus is enigmatic and provisional.

[Scyphostegia+[Prockieae+Saliceae+Homalieae+Scolopieae+Oncobeae]]

Lamina with small vein proceeding into tooth and expanding. Salicoid teeth present (tooth apex spherical and variously coloured gland or stout hair).

Scyphostegieae (Hutch.) Zmarzty in M. W. Chase et al. in Kew Bull. 57: 170. 2002

2/2. Scyphostegia (1; S. borneensis; northern and central Borneo), Dianyuea (1; D. turbinata; Yunnan). – Dioecious trees. Phellogen superficial. Vessel elements in radial multiples, with usually simple (rarely scalariform) perforation plates; lateral pits alternate. Fibres with thick walls and large bordered pits. Wood rays usually uniseriate, heterocellular. Axial parenchyma almost absent. Leaves distichous. Petiole vascular bundle transection annular; adaxial xylem and phloem developing into inverted adaxial plate of vascular tissue. Stipules small, caducous. Stomata paracytic. Inflorescence terminal, panicle. Bracts tubular, overlapping. Pedicel not articulated. Hypogyny. Perianth possibly consisting of three sepals alternating with three petals. Nectariferous lobes opposite three “antepetalous” stamens. Filaments connate. Anthers extrorse. Pollen grains tricolpate? Pistil composed of eight to 13 connate carpels. Ovary unilocular, septated near apex. Style absent. Stigma discoid, with eight to 13 rays and central orifice. Placentation basal. Ovules numerous per carpel. Micropyle bistomal (or exostomal?). Outer integument two or three cell layers thick. Inner integument three or four cell layers thick. Nucellar cap present, persistent. Fruit a fleshy capsule with lignified commissural valves. Aril formed from funicle and outer integument. Exotegmen fibrous. Perisperm very thin. Endosperm sparse. x = 9.

[Prockieae+Saliceae+Homalieae+Scolopieae+Oncobeae]

Stamens centrifugally developing. Pistil composed of two to five (to 13) connate carpels. Megagametophyte protruding through micropyle. Benzoylated glycosides present. Cyanogenic glycosides usually absent.

Prockieae Endl., Gen. Plant.: 918. Nov 1839

9/67. Banara (34; tropical America), Hasseltia (3; H. allenii, H. floribunda, H. guatemalensis; southern Mexico to Brazil and Bolivia), Hasseltiopsis (1; H. dioica; Central America), Azara (9; temperate and subtropical South America), Neosprucea (9; Panamá, northern South America), Pineda (2; P. incana, P. ovata; the Andes in Ecuador and Peru), Prockia (5; P. costaricensis, P. crucis, P. flava, P. krusei, P. pentamera; southern Mexico to northern Argentina), Abatia (10; mountain regions in tropical America), Aphaerema (1; A. spicata; southern Brazil). – Tropical America, temperate and subtropical South America. Nodes in some species of Azara bilacunar with two leaf traces. Opposite leaves present in Abatia. Tepals in Abatia with valvate aestivation, connate at base. Nectary absent in Abatia. Cyanogenic glycosides present in Banara.

Saliceae Rchb., Fl. Germ. Excurs. 1(2): 165. Jan-Apr 1831 [‘Salicinae’]

12/550–555. Populus (35–40; temperate regions on the Northern Hemisphere, on species, P. ilicifolia, in East Africa), Salix (c 450; temperate and arctic-alpine regions on the Northern Hemisphere, few species on the Southern Hemisphere), Bennettiodendron (7; B. brevipes, B. cordatum, B. lanceolatum, B. leprosipes, B. longipes, B. macrophyllum, B. subracemosum; southern China, tropical Asia), Olmediella (1; O. betschleriana; Central America), Idesia (1; I. polycarpa; China, Japan), Macrohasseltia (1; M. macroterantha; Central America), Carrierea (4; C. calycina, C. dunniana, C. rehderiana, C. vieillardii; southern and southwestern China, Southeast Asia), Itoa (1; I. orientalis; southern China, tropical Asia), Lasiochlamys (11; New Caledonia), Ludia (27; tropical East Africa, Madagascar, the Mascarene Islands), Poliothyrsis (1; P. sinensis; China), Tisonia (14; Madagascar). – Temperate and arctic-alpine regions on the Northern Hemisphere, tropical East Africa, Madagascar, the Mascarene Islands, East and tropical Asia, New Caledonia, Central America, few species on the Southern Hemisphere. Cuticular wax crystalloids as rosettes of platelets (Fabales type) or cuticular waxes absent. Inflorescence often catkin. Perianth highly reduced (petals often absent). Pollen grains in Populus inaperturate. Pistil composed of two collateral connate carpels. Ovules usually unitegmic. Micropyle in Idesia exostomal. Long placental hairs present at seed base. Tegmen impermanent. Endosperm absent in Salix. Embryo with chlorophyll. – The Idesia clade and the Salix clade appear to be very closely allied. They have salicoid leaf teeth, similar or identical phenolic compounds (e.g. salicin), and similar or identical parasites (rusts and insect larvae). Populus, Salix, Banara and Idesia with foliar extrafloral nectaries. Chosenia is nested in Salix, according to Leskinen & Alström-Rapaport (1999).

Homalieae Dumort., Anal. Fam. Plant.: 18. 1829 [‘Homalineae’]

8/c 200. Bartholomaea (3; B. mollis, B. paniculata, B. sessiliflora; Central America), Bivinia (1; B. jalbertii; tropical East Africa, Madagascar), Byrsanthus (1; B. brownii; tropical West Africa), Calantica (10; Madagascar), Dissomeria (2; D. crenata, D. glanduligera; tropical Africa), Homalium (c 180; tropical and subtropical regions on both hemispheres), Neopringlea (3; N. integrifolia, N. trinervia, N. viscosa; southern Mexico, Guatemala), Bembicia (1; B. axillaris; Madagascar). – Tropical and subtropical regions on both hemispheres. Epigyny present in Homalium.

Scolopieae Warb. in Engler et Prantl, Nat. Pflanzenfam. III, 6a: 13. 28 Dec 1893

5/43. Hemiscolopia (1; H. trimera; tropical Asia), Mocquerysia (2; M. coerulipennis, M. multiflora; tropical Africa), Phyllobotryon (2; P. bracteatum, P. paradoxum; tropical Africa), Pseudoscolopia (1; P. polyantha; South Africa), Scolopia (37; tropical regions in the Old World east to northeastern Queensland). – Tropical regions in the Old World.

Oncobeae Benth. in Proc. Linn. Soc., Bot. 5(Suppl. 2): 77. 1861

6/115–120. Oncoba (c 30; tropical and southern Africa, the Arabian Peninsula), Flacourtia (22; tropical and southern Africa, Madagascar, Southeast Asia, Malesia to Fiji), Dovyalis (17; tropical and southern Africa, Madagascar, tropical Asia from Sri Lanka to New Guinea), Trimeria (2; T. grandifolia, T. trinervis; tropical East Africa to southern Africa), Pleuranthodendron (1; P. lindenii; southern Mexico to Brazil), Xylosma (c 45?; East and Southeast Asia, Malesia to New Guinea, eastern Queensland, New Caledonia, Vanuatu, Polynesia, Guam, Central America, the West Indies, northern South America). – Pantropical. Micropyle in Oncoba endostomal. Testa in Oncoba sarcotesta. Endotesta in Oncoba palisade. – Oncoba resembles some species in Achariaceae-Lindackerieae, although they differ in, i.a., leaf teeth type, stamen initiation and phytochemistry.

Cladogram of Salicaceae (one of 709 equally most parsimonious trees of Malpighiales) based on DNA sequence data (Sosa & al. 2003). Lunania is sister to Casearia, Laetia and Samyda, according to Shang & al. (2017), and Hasseltia is sister to a clade comprising, e.g., Homalium, Xylosma, Scolopia, Oncoba, and Flacourtia.

de Jussieu in V. V. D. d’Orbigny, Dict. Univ. Hist. Nat. 12: 670. 7 Jul 1849, nom. cons.

Distribution Madagascar, West Malesia, Central America, tropical South America.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements usually with simple (rarely also scalariform) perforation plates; lateral pits almost absent, alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements tracheids and fibre tracheids (libriform fibres and intraxylary phloem absent) with bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, diffuse-in-aggregates, or banded, or paratracheal scanty, aliform, vasicentric, or banded (wood ray and axial parenchyma cell walls with helical thickenings). Wood in at least one species fluorescent. Intraxylary phloem absent. Sieve tube plastids Ss type. Nodes 1:1, unilacunar with one leaf trace; lateral bundles split. Branched sclereids present. Silica bodies absent. Rhomboid or irregularly shaped calciumoxalate crystals frequent.

Trichomes Hairs unicellular or multicellular, simple or T-shaped; glands stalked or unstalked.

Leaves Usually opposite (in Trigoniastrum alternate, spiral or distichous), simple, entire, with ? ptyxis. Stipules interpetiolar (when leaves opposite), caducous, often connate; leaf sheath absent. Petiole articulated. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata paracytic. Inconspicuous pilose domatia present on leaves in Isidodendron. Cuticular wax crystalloids? Epidermis and hypodermis with or without mucilaginous idioblasts. Leaf margin entire. Leaf margin and abaxial side of bracts in Trigoniastrum with impressed glands.

Inflorescence Terminal or axillary, panicle or thyrse (in Isidodendron raceme). Floral prophylls (bracteoles) two or three.

Flowers Obliquely zygomorphic. Hypanthium-like structure (“floral cup”) short or absent. Half epigyny. Sepals five, with imbricate quincuncial aestivation, unequal in size, connate below; median sepal adaxial. Petals three or five, usually with imbricate quincuncial or contorted (rarely valvate) aestivation, free; adaxial inner petal forming spurred or at base saccate velum together with one or two sepals; plicae of abaxial outer and abaxial lateral two petals forming often saccate carina or abaxial petals saccate; two lateral petals forming spatulate alae. One to three nectariferous glands, each up to trilobate, present at base of adaxial side of velum (large adaxial inner petal) and usually adjacent to slit on staminal tube, adnate to stamens and staminodia (nectariferous disc absent in Isidodendron, nectariferous glands here present at staminodial bases).

Androecium Five to eight (to 13) anterior four to nine stamens fertile and one to six lateral and/or four to six rudimentary posterior stamens staminodial (often scale-like), haplostemonous or diplostemonous. Filaments more or less connate into tube, slit on one side, free from tepals. Anthers dorsifixed (almost basifixed), somewhat versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often dorsally thickened? Tapetum secretory? Staminodia (one to) three to six (intrastaminal?; sometimes absent).

Pollen grains Microsporogenesis simultaneous? Pollen grains tri- to pentaporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate? infratectum, verrucate to smooth.

Gynoecium Pistil composed of three (or four) connate carpels; median carpel adaxial; carpel partially synascidiate. Ovary semi-inferior, unilocular or trilocular (or quadrilocular). Style single, simple. Stigma single, trilobate with reflexed lobes, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (parietal when ovary unilocular, with deeply intrusive placentae). Ovules one to ten (to c. 20) per carpel, anatropous, pendulous to ascending, epitropous (antitropous) to apotropous (syntropous), bitegmic, usually finally tenuinucellar by absorption of megasporangium (rarely crassinucellar). Micropyle bistomal, Z-shaped (zig-zag), or endostomal. Outer integument two or three cell layers thick. Inner integument four to six cell layers thick, entirely endothelial. Obturator consisting of unicellular funicular hairs or papillae. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Usually a septicidal capsule with carpels internally dehiscing and with persistent central fibre strands (in Humbertiodendron and Trigoniastrum a three-winged samaroid).

Seeds Aril absent. Testa sometimes winged, often with long lignified hairs. Exotestal cells with thickened outer walls. Endotesta? Tegmen often multiplicative. Exotegmen? Endotegmic cells with tannins and somewhat thickened walls. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, in Trigonia with chlorophyll. Cotyledons two, large, thin, flat. Hypocotyl elongate. Germination phanerocotylar.

Phytochemistry Virtually unknown. Tannins present. Aluminium not accumulated.

Piriquetaceae Martinov, Tekhno-Bot. Slovar: 484. 3 Aug 1820 [’Piriquettae’]; Turnerales Link, Handbuch 2: 47. 4-11 Jul 1829 [’Turneraceae’]

Distribution Tropical, subtropical and warm-temperate regions in Mexico, Central and South America, the West Indies and Africa south of Sahara, Madagascar, Rodriguez.

Habit Bisexual, usually perennial herbs or evergreen shrubs (rarely trees). Often evil-smelling.

Vegetative anatomy Phellogen ab initio superficial. Cortical vascular bundles abundant. Medulla usually parenchymatous, sometimes with sclereids. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits? Imperforate tracheary xylem elements libriform fibres with thick walls and small bordered pits, non-septate? (also vasicentric tracheids). Wood rays uniseriate or multiseriate?, heterocellular. Axial parenchyma apotracheal diffuse? Sieve tube plastids Ss type; sieve tubes with non-dispersive protein bodies? Nodes? Crystalliferous and tanniniferous cells frequent; parenchyma cells sometimes with calciumoxalate crystals and starch grains.

Trichomes Hairs unicellular or multicellular, usually uniseriate, simple, sometimes multi-armed or stellate; glandular hairs excreting nasty-smelling substance often present.

Leaves Alternate (spiral), usually simple (simetimes pinnately compound), entire, sometimes ericoid, usually with conduplicate (sometimes revolute) ptyxis. Stipules usually small or absent (present in Erblichia and some species of Turnera); leaf sheath absent. Stipules and/or prophylls and leaf primordia with colleters. Petiole vascular bundle transection? Distal part of petiole or leaf base often with extrafloral nectaries. Venation pinnate, with coarse veins; vein proceeding into transparent and caducous tooth apex? Stomata anomocytic, paracytic or anisocytic. Cuticular wax crystalloids? Lamina sometimes gland-dotted. Epidermis with or without mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts with calciumoxalate druses. Leaf margin serrate or crenate (teeth sometimes glanduliferous).

Inflorescence Flowers usually solitary axillary, sometimes in terminal or axillary raceme (flowers in Piriqueta and several species of Turnera epiphyllous). Floral prophylls (bracteoles) often large.

Flowers Actinomorphic. Hypogyny or half epigyny. Sepals five, with imbricate quincuncial aestivation, usually connate into caducous tube (in Erblichia and Mathurina almost free). Petals five, with contorted aestivation, clawed, early withering and caducous, usually with claws adnate to calyx and together with this forming tubular, campanulate or infundibuliform structure (hypanthium; in Erblichia and Mathurina free); in orifice sometimes five nectariferous glands, fringed corona or five lobes between corona and stamens (nectaries sometimes present in perianth tube, in Mathurina and Stapfiella on sepals, in some species of Turnera on stamens or as five nectariferous pockets). Disc extrastaminal, annular, or absent. Heterostyly sometimes present.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments usually free from each other, usually free from tepals (sometimes adnate at base to petals; filaments in Erblichia and Hyalocalyx adnate at base to calyx). Anthers usually dorsifixed (in Erblichia and some species of Turnera almost basifixed), sometimes versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of (two or) three connate carpels. Ovary superior to semi-inferior, unilocular. Stylodia (two or) three, filiform, often bifid, free. Stigmas concave, commissural, fringed and bristle-like, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation usually parietal (in Stapfiella basal). Ovules one (Stapfiella) to more than 100 per ovary (Stapfiella with one basal ovule), anatropous, ascending, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Hypostase present. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit A loculicidal capsule, usually dehiscing from apex (rarely laterally), sometimes with persistent perianth tube.

Seeds Aril or elaiosome (in Mathurina fimbriate) often present. Operculum present. Testa hard, non-multiplicative. Exotestal cells arranged in rows. Exotegmen palisade, with sclereids. Endotegmen persistent? Perisperm not developed. Endosperm copious, fleshy, oily. Embryo straight, well differentiated, chlorophyll? Cotyledons two, plano-convex. Germination phanerocotylar.

Phytochemistry Insufficiently known. Luteolin, ethereal oils, tannins, alkaloids, and cyclopentenoid cyanogenic glycosides and/or cyclopentenylic fatty acids present. Flavonols, ellagic acid, and proanthocyanidins not found.

Violales Vent. ex Bercht. et J. Presl, Přir. Rostlin: 220. Jan-Apr 1820 [‘Violaceae’]; Ionidiaceae Mert. et W. J. Koch in J. C. Röhlings, Deutschl. Fl., ed. 3, 1(1): 244, 260. Jan-Mai 1823 [’Jonidien’, ’Jonidiae’], nom. illeg.; Ionidiales Vent. in C. F. P. von Martius, Consp. Regn. Veg.: 50. Sep-Oct 1835 [‘Jonidieae’], nom. illeg.; Violopsida Brongn., Enum. Plant. Mus. Paris: xxiv, 88. 12 Aug 1843 [’Violineae’]; Leoniaceae A. DC. in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 8: 668. med Mar 1844; Alsodeiaceae J. Agardh, Theoria Syst. Plant.: 197. Apr-Sep 1858 [‘Alsodineae’]; Violanae R. Dahlgren ex Reveal in Novon 2: 237. 13 Oct 1992

Habit Usually bisexual (in Melicytus dioecious), perennial or annual herbs, evergreen or deciduous shrubs or trees (rarely lianas).

Vegetative anatomy Phellogen superficial. Primary vascular tissue cylinder without separate vascular bundles, or cylinder of bundles. Cambium in Viola storied. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, opposite or scalariform, usually simple (sometimes bordered) pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular (sometimes absent). Axial parenchyma rare (paratracheal scanty) or absent. Sieve tube plastids S type; sieve tubes with non-dispersive protein bodies? Endodermis in Viola cunninghamii consisting of large cells with suberised walls. Nodes usually 3:3, trilacunar with three leaf traces (in Fusispermum 5:5, quinquelacunar with five traces). Prismatic calciumoxalate crystals and druses frequent.

Trichomes Hairs unicellular or multicellular, uniseriate, simple; glandular hairs sometimes present.

Leaves Usually alternate (spiral or distichous; in some species of Hybanthus and Rinorea opposite), simple, entire or lobed, with often involute ptyxis. Stipules sometimes petiolar, foliaceous, often lobed; leaf sheath absent. Colleters usually present. Petiole vascular bundle transection usually arcuate. Venation pinnate or palmate. Stomata paracytic or anomocytic (anisocytic?), often on adaxial side of lamina only. Cuticular wax crystalloids? Domatia as pits or hair tufts. Epidermis often with mucilaginous idioblasts. Secretory cavities absent. Leaf margin serrate or entire.

Inflorescence Terminal or axillary, panicle, raceme-like or capitate, or flowers solitary axillary.

Flowers Zygomorphic (in, i.a., Melicytus almost actinomorphic). Pedicel articulated. Hypogyny. Sepals five, with median sepal adaxial, with imbricate quincuncial or open aestivation, often persistent, free or connate at base, sometimes with basal appendages. Petals five, usually with apotact (in Fusispermum convolute, in Leonia and some species of Gloeospermum imbricate quincuncial) aestivation, free, abaxial (anterior) petal often with nectariferous spur directed backwards. Nectaries present on lower part of stamens or absent. Disc usually absent (in Fusispermum fleshy, annular, quinquelobate).

Androecium Stamens usually five (in Leonia triandra three), antesepalous, alternipetalous (in Fusispermum with very small fringed apical scales). Filaments usually free (sometimes connate at base into tube; in Fusispermum adnate with indentations to inner surface of petals), free from tepals; all or two abaxial stamens often with abaxial nectariferous appendage in lower part, or with prolonged connective (anthers of Leonia without dorsal connective scales). Anthers basifixed to laterofixed, usually connivent forming ring, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits; thecae in Fusispermum cordate/trapezoid); connective often prolonged into membranous scale-like appendage. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–5)-colporate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or microreticulate.

Gynoecium Pistil composed of (two or) three (in Leonia five, antesepalous) connate carpels; median carpel abaxial. Ovary superior, unilocular. Style single, simple, straight or curved, or stylodia (two or) three (in Leonia five), free (absent in Melicytus). Stigma subcapitate or lobate, often asymmetrical (stigmas in Melicytus three to five), papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal. Ovules one to numerous per ovary, anatropous, bitegmic, crassinucellar. Micropyle usually bistomal, Z-shaped (zig-zag; rarely endostomal). Outer integument two to four cell layers thick. Inner integument approx. three cell layers thick. Hypostase present. Parietal tissue two or three cell layers thick. Nucellar cap approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids rarely with a filiform apparatus. Antipodal cells usually ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit Usually an explosive loculicidal capsule (in Leonia a nut; in Melicytus berry), often with persistent calyx. Maturing seeds in Anchietea and Decorsella exposed on open carpels.

Seeds Seeds often with aril and/or carunculus (in Viola sometimes with elaiosome). Seed coat endotestal-exotegmic, in Gloeospermum and Leonia mucilaginous, subglobose. Testa in Agatea, Anchietea and Corynostylis winged. Exotesta subpalisade to tabular, sometimes with thickened cell walls. Mesotesta sometimes sclerenchymatous. Endotesta cells usually with calciumoxalate crystals. Fibre layer one to three cell layers thick. Exotegmen fibrous; exotegmic cells tracheidal, with thickened lignified walls. Endotegmen persistent, usually crystalliferous. Perisperm not developed. Endosperm usually copious, oily (in Gloeospermum absent). Embryo usually large, straight, with chlorophyll (at least in Viola). Cotyledons two, flat. Germination phanerocotylar.

DNA Plastid gene rps16 absent in at least two species of Viola. Mitochondrial coxI intron present.

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin), alkaloids, saponins, and cyclotide proteins present. Ellagic acid, tannins, proanthocyanidins and cyanogenic compounds not found. Aluminium accumulated in some species.

Use Ornamental plants, medicinal plants, perfumes, flavouring and confection (Viola odorata), timber.

Systematics Violaceae are sister-group to the clade [Malesherbiaceae+[Passifloraceae+Turneraceae]].

Fusispermum is sister to the remaining Violaceae.

Fusispermoideae Hekking in Proc. Kon. Ned. Akad. Wetensch., ser. C, 7: 128. 18 Jun 1984

1/3. Fusispermum (3; F. laxiflorum, F. minutiflorum, F. rubrolignosum; Panamá, Colombia, Peru). – Medullary cells thin-walled. Nodes 5:5, quinquelacunar with five leaf traces. Petiole with elliptical medullary bundle. Phloem internal. Petals with contorted aestivation. Disc annular, quinquelobate, carnose, with lobes alternating with stamens. Filaments adnate to inner surface at indentations. Thecae cordate/trapezoid, confluent at apex?; connective as short paired fimbriate ventral apical scales. Fruit a loculicidal capsule. Seeds elongate, longitudinally winged. Exotegmen moderately developed?, with slightly elongate cells. n = ?

Violoideae Beilschm. in Flora 16(Beibl. 7): 88. 14 Jun 1833 [‘Violieae’]

c 30/900–1.000. ‘Rinorea’ crenata group (3; R. apiculata, R. crenata; southern Central America, tropical South America); Rinorea (225–275; tropical regions on both hemispheres); Viola (580–620; temperate and alpine regions on both hemispheres, the Andes, tropical mountains), Allexis (3; A. cauliflora, A. obanensis, A. zygomorpha; Ghana, Nigeria, Cameroon), Noisettia (1; N. orchidiflora; Guyana, Brazil, Peru), Schweiggeria (1; S. fruticosa; eastern Brazil); Rinoreocarpus (1; R. ulei; Amazonia); Decorsella (2; D. arborea, D. paradoxa; tropical tropical West Africa), Paypayrola (8; southern Central America, tropical South America; incl. Hekkingia?), Hekkingia (1; H. bordenavei; French Guiana, northern Brazil; in Paypayrola?); Leonia (4; L. crassa, L. cymosa, L. glycycarpa, L. triandra; southern Central America, tropical South America; non-monophyletic?), Gloeospermum (c 12; Central America, northern South America), Amphirrhox (1; A. longifolia; Central America, northern South America), Pigea (c 10; southern Australia, New Caledonia, islands in southern Pacific), Hybanthus (3; H. concolor, H. havanensis, H. yucatanensis; eastern North America, Mexico, the West Indies), Mayanaea (1; M. caudata; Guatemala), Orthion (5–6; O. guatemalense, O. malpighiifolium, O. montanum, O. oblanceolatum, O. subsessile, O. veracruzense; Central America, northern South America); ‘Rinorea’ virgata group (1; R. virgata; southern India, Sri Lanka, the Andaman Islands, Southeast Asia), ‘Hybanthus’ enneaspermus group (c 25; Africa, the Arabian Peninsula, India, northern Australia), ‘Hybanthus’ fruticulosus group (3; H. fruticulosus, H. serrulatus; Mexico, Central America), ‘Hybanthus’ thiemei group (4–6; H. galeotii, H. nanus, H. thiemei; Central America, the West Indies, northern South America), Ixchelia (2; I. mexicanus, I. uxpanapana; Mexico, Central America), Isodendrion (4; I. hosakae, I. laurifolium, I. longifolium, I. pyrifolium; the Hawaiian Islands), Pombalia (c 65; southwestern United States, Central America, the West Indies, tropical South America), Melicytus (c 10; southeastern South Australia, Victoria, eastern New South Wales, Tasmania, Solomon Islands, Norfolk Island, New Zealand, Fiji, the Hawaiian Islands), ‘Hybanthus’ guanacastensis group (2; H. denticulatus, H. guanacastensis; Mexico, Central America), Anchietea (9–10; tropical South America), Hybanthopsis (1; H. bahiensis; eastern Brazil), Calyptrion (4; C. arboreum, C. carthagenense, C. pubescens, C. volubile; Central America, tropical South America), Agatea (c 8; New Guinea, New Caledonia, islands in southern Pacific). – Cosmopolitan. Petals often with quincuncial aestivation. Nectary lobes opposite stamens; nectaries usually separately adnate to filament bases. Anther thecae sometimes horizontal; connective lobate, as long as and wider than anther, free, with apex not or very slightly bilobate and margin entire or erose, usually covering thecae from abaxial side. Stigmatic head subcapitate; receptive small. – Rinorea apiculata and R. crenata form a sister-group to the remaining Violoideae (Wahlert 2014). Rinorea has nectary shaped as thick semicircular lobes at filament bases, thecae visible from abaxial side, subapical anther connective, style widening subapically, and one ovule per carpel.

Cladogram (simplified) of Violaceae based on DNA sequence data (Tokuoka 2008). Ixchelia is sister to [Isodendrion+Pombalia], according to Wahlert & al. (2015).

Abdul-Salim K. 2002. Systematics and biology of Symphonia L. f. (Clusiaceae). – Ph.D. diss., Harvard University, Cambridge, Massachusetts.

Abou-Shoer M, Suwqanborirux K, Habib AAM, Chang CJ, Cassady JM. 1993. Xanthones and vismiones from Psorospermum febrifugum. – Phytochemistry Nov 1993. v.34 (5).

Abreu V, Bove C, Philbrick C, Mendonça C, Gonçalves-Esteves V. 2012. Pollen morphology of the aquatic Brazilian endemic genus Castelnavia Tul. & Wedd. (Podostemaceae). – Plant Syst. Evol. 298: 1455-1461.

Achoundong G. 2003. Novitates Gabonenses 45. Une nouvelle espèce de Rinorea (Violaceae) du Gabon. – Adansonia, sér. III, 25: 211-214.

Achoundong G, Bakker FT. 2006. Deux nouvelles espèces de Rinorea, série Ilicifoliae (Violaceae) du Cameroun. – Adansonia, sér. III, 28: 129-136.

Achoundong G, Bos JJ. 2001. Deux espèces nouvelles de Rinorea (Violaceae) du Congo et du Gabon. – Adansonia, sér. III, 23: 155-159.

Achoundong G, Cheek M. 2003. Two new species of Rinorea (Violaceae) from Western Cameroon. – Kew Bull. 58: 957-964.

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”The COM clade”

MALPIGHIALES Juss. ex Bercht. et J. Presl

ACHARIACEAE Harms

BALANOPACEAE Benth. et J. D. Hooker

BONNETIACEAE (Bartl.) L. Beauvis. ex Nakai

CALOPHYLLACEAE J. Agardh

CARYOCARACEAE Voigt

CENTROPLACACEAE (Radcl.-Sm.) Doweld et Reveal

CHRYSOBALANACEAE R. Br.

CLUSIACEAE Lindl.

CTENOLOPHONACEAE (H. Winkl.) Exell et Mendonça

DICHAPETALACEAE Baill.

ELATINACEAE Dumort.

ERYTHROXYLACEAE Kunth

EUPHORBIACEAE Juss.

EUPHRONIACEAE Marc.-Berti

GOUPIACEAE Miers

HUMIRIACEAE A. Juss.

HYPERICACEAE Juss.

IRVINGIACEAE (Engl.) Exell et Mendonça

IXONANTHACEAE Planch. ex Miq.

LACISTEMATACEAE Mart.

LINACEAE DC. ex Perleb

LOPHOPYXIDACEAE (Engl.) H. Pfeiffer

MALESHERBIACEAE D. Don

MALPIGHIACEAE Juss.

MEDUSAGYNACEAE Engler et Gilg, nom. cons.

OCHNACEAE DC.

PANDACEAE Pierre ex Engl. et Gilg

PASSIFLORACEAE Juss. ex Roussel

PERACEAE (Baill.) Benth. ex Klotsch

PHYLLANTHACEAE Martinov

PICRODENDRACEAE Small

PODOSTEMACEAE Rich. ex Kunth

PUTRANJIVACEAE (Endl.) Meisn.

QUIINACEAE Choisy ex Engl.

RAFFLESIACEAE Dumort.

RHIZOPHORACEAE Pers.

SALICACEAE Mirb.

TRIGONIACEAE A. Juss.

TURNERACEAE Kunth ex DC.

VIOLACEAE Batsch