MYRTANAE Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 295. 4 Feb 1967

Myrtopsida Bartl., Ord. Nat. Plant.: 225, 326. Sep 1830 [’Myrtinae’]; Myrtidae J. H. Schaffn., Ohio Naturalist [’Myrtiflorae’] 11: 416. Dec 1911; Geranianae Thorne ex Reveal in Novon 2: 236. 13 Oct 1992

[Geraniales+Myrtales]


MYRTALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 233. Jan-Apr 1820 [‘Myrtaceae’]

Fossils Esgueiria includes fossilized reproductive organs which have been attributed to Myrtaceae. Esgueiria, from the Campanian to the Maastrichtian of Portugal and Japan, is represented by epigynous flowers with a persistent calyx of five sepals, five petals, 3+5 stamens, tricolpate pollen grains, an annular (nectariferous) disc, three stylodia and up to six apical anatropous ovules in the unilocular ovary; the calyx and the ovary possess abundant simple hairs and peltate multicellular glands.

Habit Usually bisexual (rarely andromonoecious, polygamomonoecious, dioecious or androdioecious, polygamodioecious), usually evergreen (rarely deciduous) trees or shrubs, or perennial or annual herbs (sometimes lianas). Young stems and branches often quadrangular in transverse section.

Vegetative anatomy Phellogen ab initio subepidermal, pericyclic or cortical. Polyderm often present. Secondary lateral growth normal, anomalous (from cylindrical cambium) or absent. Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits usually alternate (sometimes opposite), simple or bordered pits. Vestured pits usually present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres, with simple or bordered pits, septate or non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, often banded, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, reticulate, scalariform, unilateral, vasicentric or banded. Intraxylary phloem usually present. Interxylary phloem sometimes present. Secondary phloem in young stems usually stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids Ss type. Nodes 1:1 or 1:3, unilacunar with one or three leaf traces, or 3:3, trilacunar with three traces. Schizolysigenous secretory cavities with ethereal oils. Heartwood often with gum-like substances. Silica bodies sometimes present. Calciumoxalate as prismatic, rhomboidal or acicular crystals, styloids, or crystal sand often frequent (raphides usually absent, sometimes frequent).

Trichomes Hairs unicellular, bicellular or multicellular, uniseriate or multiseriate, simple or branched, furcate, stellate, dendritic, peltate, lepidote or vesicular, or absent; glands or glandular hairs (bristle-like or lepidote) often present.

Leaves Usually opposite (sometimes verticillate or alternate), simple, entire, often coriaceous, with conduplicate, supervolute, revolute, involute or flat ptyxis. Stipules usually rudimentary non-vascularized or absent (sometimes hair-like, colleter-like or glandular, rarely paired or few and intrapetiolar); leaf sheath absent. Colleters often frequent. Petiole vascular bundle transection arcuate or annular. Venation pinnate or palmate, brochidodromous, eucamptodromous, craspedodromous, parallelodromous, palmate-parallel or acrodromous (leaves sometimes one-veined). Stomata usually anomocytic (sometimes anisocytic, paracytic, polycytic, diacytic, tetracytic, cyclocytic or anomocyclocytic). Cuticular wax crystalloids sometimes as rosettes, often absent. Domatia as pits, pockets or hair tufts (acarodomatia or myrmecodomatia sometimes present). Lamina with gland-dots and schizogenous secretory cavities with ethereal oils. Epidermis with or without mucilaginous idioblasts. Mesophyll often with sclerenchymatous idioblasts containing sclereids of different types and with calciumoxalate as single prismatic crystals, styloids, druses or crystal sand. Leaf margin usually entire (rarely serrate or dentate).

Inflorescence Terminal or axillary, panicle or thyrse, simple or branched racemose, spicate, umbellate or capitate, or corymb (flowers sometimes solitary). Floral prophylls (bracteoles) sometimes absent.

Flowers Usually actinomorphic (sometimes zygomorphic, rarely asymmetrical). Hypanthium present, usually elongate. Usually epigyny or half epigyny (rarely hypogyny). Sepals (two to) four to eight (to 16), with imbricate, valvate, contorted or open (rarely cochlear) aestivation, usually free (sometimes connate into caducous calyptra). Petals (two to) four to eight (to numerous), with imbricate, valvate, plicate or contorted aestivation, often clawed, free or connate at base (sometimes forming caducous calyptra or absent). Nectariferous disc annular or unilateral at ovary tip, ovary base, along elongated hypanthium, or intrastaminal, or nectaries on adaxial side of hypanthium and disc intrastaminal on top of ovary, or nectary and disc absent.

Androecium Stamens (one to) six to ten in (one or) two whorls, or c. 20 to more than 150 in one or several whorls, alternisepalous or antesepalous, alternipetalous or antepetalous. Filaments free or connate at base into four or five fascicles, or connate into tube, usually free from tepals (rarely epipetalous), inserted on hypanthium, often incurved in bud. Anthers basifixed or dorsifixed, usually versatile, usually tetrasporangiate (rarely monosporangiate, disporangiate or trisporangiate), introrse, latrorse or extrorse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by pores). Tapetum secretory. Staminodia usually absent (rarely four to ten extrastaminal, or four or five alternating with fertile stamens; female flowers rarely with staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colpor(oid)ate, (2–)3(–6)-colpate or (2–)3(–6)-por(or)ate, often heterocolpate (with pseudocolpi alternating with apertures), usually shed as monads (occasionally as tetrads or polyads), bicellular at dispersal. Exine tectate or semitectate, with columellate (rarely acolumellate) infratectum, perforate, punctate, reticulate, striate, rugulate, verrucate, echinate, scabrate, granulate or smooth.

Gynoecium Pistil composed of two to five (to 16) connate carpels. Ovary usually inferior or semi-inferior (rarely superior), unilocular to quinquelocular (rarely up to 16-locular). Style single, sunken, usually elongate, persistent, entire or branched above. Stigma usually single, simple, capitate, or tri- or quadrilobate (stigmas then three or four, punctate), papillate, Dry or Wet type. Male flowers sometimes with pistillodium.

Ovules Placentation usually axile (sometimes basal, parietal or free central, rarely apical-axile). Ovules (one or) two to c. 30 (to more than 150) per carpel, anatropous, hemianatropous or campylotropous (rarely orthotropous), ascending or horizontal (rarely pendulous), apotropous or epitropous, usually bitegmic (rarely unitegmic), crassinucellar. Micropyle bistomal (often Z-shaped) or endostomal. Funicular obturator sometimes present. Nucellar cap often present (nucellar beak rarely present). Archespore sometimes multicellular. Megagametophyte usually monosporous, Polygonum type (sometimes quadrinucleate, Oenothera type, rarely disporous, Allium type, or tetrasporous, 16-nucleate, Penaea type). Synergids often with a filiform apparatus. Antipodal cells usually ephemeral, sometimes not developed (nuclei early degenerating). Endosperm development ab initio nuclear. Endosperm haustoria absent? Embryogenesis usually onagrad (sometimes asterad or solanad).

Fruit Usually a loculicidal (sometimes septicidal or irregularly dehiscing) capsule, a berry or nutlike fruit (rarely a denticidal capsule, pyxidium, drupe, or schizocarp), often with calyx persistent and accrescent.

Seeds Aril absent. Funicular elaiosome rarely present. Testa sometimes winged. Operculum, formed by hilum, often present. Seed coat testal. Testa often multilayered, sometimes multiplicative, sometimes with crystalliferous layers. Exotesta sometimes palisade to cuboidal and lignified. Mesotesta sometimes sclerotic. Endotesta often crystalliferous, sometimes collapsed. Tegmen often crushed or exotegmen fibrous or tracheidal. Endotegmen usually crushed (rarely fibrous). Perisperm usually not developed. Endosperm usually absent (sometimes scarce, oily). Embryo small, straight, curved or spirally twisted, well differentiated, with or without chlorophyll. Cotyledons (one or) two (to five). Germination phanerocotylar or cryptocotylar.

Cytology x = (5–)7–15 (rarely higher)

DNA Plastid gene infA lost/defunct. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), acylated anthocyanins, flavonoid sulfates, flavonoid mono- and diglycosides, cyanidin, delphinidin, ethereal oils (triterpenes, etc.), ursolic acid, ellagic and gallic acids, methylated ellagic acids, galloyl- and ellagitannins (e.g. cuphiin), non-hydrolyzable tannins, polyhydroxyalkaloids (usually frequent), quinolizidine alkaloids, triterpene saponins, phenylalanine-derived cyanogenic compounds (prunasin etc.), anthraquinones, and polyacetate-derived arthroquinones present. Raffinose and stachyose in phloem exudates. Aluminium or oxalate accumulated in some species.

Systematics Myrtales are sister-group to Geraniales (Wang & al. 2009).

A plausible topology is [Combretaceae+[Lythraceae+Onagraceae]+[[Vochysiaceae+ Myrtaceae]+[[Melastomataceae+Memecylaceae]+[Crypteroniaceae+[Alzateaceae+[Rhyncho-calycaceae+[Oliniaceae+Penaeaceae]]]]]]].

Lythraceae and Onagraceae share the following potential synapomorphies (Stevens 2001 onwards): vessels present in groups in wood; fibres with simple or at most minutely bordered pits; petiole vascular bundle transection arcuate; pollen grains at anthesis sometimes starchy; presence of hypostase; megasporocytes several; megasporangium with starch grains; sepals persistent in fruit; exotegmen fibrous; x = 8; tannins often not abundant; and soluble oxalate accumulated.

The clade [[Vochysiaceae+Myrtaceae]+[[Melastomataceae+Memecylaceae]+[Cryptero-niaceae+[Alzateaceae+[Rhynchocalycaceae+[Oliniaceae+Penaeaceae]]]]]] is characterized by the potential synapomorphy: inflorescence with at least branches cymose.

Myrtaceae and Vochysiaceae share the characters (Stevens 2001 onwards): hairs unicellular or bicellular, simple; sepals and petals with imbricate aestivation; pollen grains syncolporate; style depressed in apex of gynoecium; and fruit a capsule.

The clade [[Melastomataceae+Memecylaceae]+[Crypteroniaceae+[Alzateaceae+[Rhyncho-calycaceae+ [Oliniaceae+Penaeaceae]]]]] has the following potential synapomorphies in common (Stevens 2001 onwards): branched or unbranched sclereids present or absent within same major clade; absence of nectary; connective abaxially much expanded; and endothecial thickening absent or atypical.

Melastomataceae and Memecylaceae share numerous potentially advanced features including (Stevens 2001 onwards): presence of internal phloem; presence of two or four distinct lateral veins arising from near base of lamina; sepals with imbricate quincuncial aestivation; petals with contorted aestivation; anthers poricidal, inverted during development, with branched vascular trace; expansion of connective below anther well developed, with various appendages; pollen grains with pseudocolpi; carpels antepetalous; stigma punctate; outer and inner integuments approx. two cell layers thick; and radicula curved.

The clade [Crypteroniaceae+[Alzateaceae+[Rhynchocalycaceae+[Oliniaceae+Penaeaceae]]]] has the following potential synapomorphies: vessel/ray and vessel/parenchyma pits half-bordered; stipules minute; stamens as many as petals, antepetalous; endothecium ephemeral; fruit a capsule; exotestal cells periclinally elongated; and endotegmen not fibrous.

The clade [Alzateaceae+[Rhynchocalycaceae+[Penaeaceae+Oliniaceae]]] is characterized by stout style. The [Rhynchocalycaceae+[Penaeaceae+Oliniaceae]] clade has: leaves with glandular apex; and x = 10. Finally, Oliniaceae and Penaeaceae share the characters (Stevens 2001 onwards): hypanthium well developed; pollen grains psilate; foot layer and tectum thick; exotegmen fibrous; absence of embryo suspensor; presence of non-hydrolyzable tannins; and aluminium not accumulated.

Cladogram of Myrtales based on DNA sequence data (Conti & al. 1997). The major clades are well supported, although the position of Combretaceae is uncertain; they are sometimes recovered as sister-group either to the remainder or to the clade [Lythraceae+Onagraceae] (e.g. Soltis & al. 2011, with low support).

ALZATEACEAE S. A. Graham

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Graham in Ann. Missouri Bot. Gard. 71: 775. 19 Apr 1985

Genera/species 1/1

Distribution The Andes from Costa Rica to Bolivia.

Fossils Unknown.

Habit Bisexual, evergreen trees or shrubs (sometimes semi-epiphytic). Young stems and branches at first quadrangular in cross-section, later terete.

Vegetative anatomy Phellogen ab initio deeply seated, close to perivascular sclerenchyma. Cortical vascular bundles present. Primary vascular tissue cylinder, without separate vascular bundles, bicollateral. Vessel elements diffuse, with simple perforation plates; lateral pits alternate?, bordered pits. Vessel/ray and vessel/parenchyma pits simple. Vestured pits present. Imperforate tracheary xylem elements thin-walled libriform fibres with simple pits (almost without bordered pits), septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal scanty (apotracheal parenchyma absent). Intraxylary phloem present. Sieve tube plastids Ss type? Nodes 3:3, trilacunar with three leaf traces. Phloem parenchyma with druses. Sclereids present. Solitary crystals absent.

Trichomes Hairs absent.

Leaves Opposite or verticillate, simple, entire, coriaceous, with ? ptyxis. Stipules small, paired or few together, intrapetiolar (axillary), or absent; leaf sheath absent. Petiole vascular bundle transection annular; petiole with wing bundles. Venation pinnate, brochidodromous. Stomata anomocytic, laterocytic or cyclocytic. Cuticular wax crystalloids? Mesophyll with sclerenchymatous idioblasts (containing sclereids) and calciumoxalate druses. Leaf margin entire.

Inflorescence Axillary, thyrse or panicle. Bracteoles absent.

Flowers Actinomorphic, small. Hypanthium short. Hypogyny (to half epigyny). Sepals five (or six), with valvate aestivation, thick, free, pointed in bud. Petals absent (sometimes rudimentary). Nectariferous disc intrastaminal, lobate.

Androecium Stamens five, alternisepalous. Filaments short, inserted at margin of nectariferous disc, free from each other and from tepals. Anthers dorsifixed, with terminal microsporangia (inserted along apex of connective), versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective wide, cordately expanded, with appendage directed backwards; endothecium ephemeral. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular at dispersal. Not heterocolpate (pseudocolpi almost absent). Exine tectate?, with columellate infratectum, psilate to verrucate-areolate.

Gynoecium Pistil composed of two connate carpels. Ovary superior (to semi-inferior), bilocular, with incomplete septum. Transseptal vascular bundles present. Style single, simple, short. Stigma capitate, Wet type? Pistillodium absent.

Ovules Placentation intrusively parietal. Ovules c. 40 to c. 60 per carpel, anatropous, horizontal, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Archespore multicellular. Megagametophyte disporous, 8-nucleate, Allium type. Endosperm development probably nuclear. Endosperm haustoria? Embryogenesis?

Fruit A flattened loculicidal capsule, with persistent calyx.

Seeds Aril absent. Seeds flattened. Testa winged, with hairpin-shaped vascular bundle. Seed coat exotestal. Exotestal cells low, with irregularly sinuate anticlinal walls. Endotesta and tegmen collapsed. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = 14

DNA

Phytochemistry Insufficiently known. Flavonol mono- and diglycosides (quercetin-3-O-glycoside, quercetin-3-O-diglycoside, etc.) and ellagic acid present. Aluminium not accumulated. Myricetin- and flavone-C-glycosides not found.

Use Unknown.

Systematics: Alzatea (1; A. verticillata; cloud forests of Costa Rica and Panamà, lower Andean slopes of Peru and Bolivia).

Alzatea is sister to the clade [Rhynchocalyx+[Olinia+Penaeaceae]].

COMBRETACEAE R. Br.

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Brown, Prodr. Fl. Nov.-Holl.: 351. 27 Mar 1810, nom. cons.

Terminaliaceae J. St.-Hil., Expos. Fam. Nat. 1: 178. Feb-Apr 1805; Combretales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 234. Jan-Apr 1820 [‘Combretaceae’]; Myrobalanaceae Juss. ex Martinov, Tekhno-Bot. Slovar: 396. 3 Aug 1820 [’Mirobolaneae’]; Bucidaceae Spreng., Syst. Veg. 2: 282. Jan-Mar 1825 [’Bucideae’]; Myrobalanales Link, Handbook 2: 440. 4-11 Jul 1829 [‘Myrobalaneae’]

Genera/species 12/495–530

Distribution Pantropical.

Fossils Terminalioxylon is fossil wood assigned to Combretaceae known from numerous Cenozoic sites in Africa, South America and tropical Asia. Pollen grains have been recorded from the Late Eocene onwards.

Habit Usually bisexual (rarely andromonoecious or dioecious), evergreen or deciduous trees, shrubs, suffrutices or lianas (Lumnitzera and Laguncularia are mangrove trees with pneumatophores). Bark often exfoliating. Sometimes xerophytic.

Vegetative anatomy Phellogen ab initio usually cortical? or epidermal (sometimes deeper). Secondary lateral growth normal or anomalous (from cylindrical cambium). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids and often libriform fibres with usually simple pits (sometimes very small bordered pits; e.g. Strephonema), septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, scalariform, reticulate, unilateral, vasicentric or banded. Tyloses sometimes frequent. Intraxylary and/or interxylary phloem present in some genera, in cross-section often distributed as reticulated islands. Sieve tube plastids Ss type. Nodes 1:1, unilacunar with one leaf trace. Sclereids present or absent. Mucilage ducts present in Terminalia. Heartwood sometimes with gum-like substances. Silica bodies present in some species. Calciumoxalate raphides absent. Prismatic crystals often frequent. Styloids or druses sometimes present.

Trichomes Hairs usually unicellular (sometimes multicellular), thick-walled, often pointed, with distinct basal segment (‘combretaceous hairs’), simple or peltate-lepidote (sometimes furcate or vesicular), often adpressed; glandular hairs often present (sometimes lepidote).

Leaves Alternate (often in two or four rows) or opposite (rarely verticillate), simple, entire, with conduplicate or supervolute (in Laguncularia revolute) ptyxis. Stipules rudimentary or absent; leaf sheath absent. Petiole vascular bundle transection arcuate to annular; wing bundles present in some species. Venation pinnate, brochidodromous, eucamptodromous or craspedodromous. Stomata usually anomocytic (in Strephonema paracytic; in Laguncularia and Lumnitzera cyclocytic). Cuticular wax crystalloids as rosettes of platelets (Fabales type). Domatia as pits, pockets or hair tufts numerous, as well as glands (lamina sometimes gland-dotted) and/or pellucid dots. Large mesophyll cells with solitary calciumoxalate druse. Mucilaginous idioblasts cells present or absent. Hydathodes sometimes present. Leaf margin entire. Paired extrafloral nectaries often present on petiole; foliar nectaries present in many species.

Inflorescence Terminal or axillary, simple or branched spicate or raceme (sometimes congested, rarely umbellate).

Flowers Usually actinomorphic (sometimes weakly zygomorphic). Hypanthium elongate. Usually epigyny (in Strephonema half epigyny). Sepals four or five (to eight), in one whorl, with imbricate or valvate aestivation, connate (in Getonia persistent and accrescent in fruit). Petals four or five, with imbricate or valvate aestivation, sometimes clawed, connate (absent in some species). Nectariferous disc present at base of upper part of hypanthium, or nectary and disc absent.

Androecium Stamens usually 4+4 or 5+5 (rarely four, five or 8+8), obdiplostemonous, alternipetalous or antepetalous. Filaments inserted on adaxial side of upper part of hypanthium (usually at two levels; one staminal whorl rarely staminodial), usually free from each other and from tepals (in some species of Terminalia adnate to petals, epipetalous). Anthers dorsifixed, usually versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, usually binucleate (rarely trinucleate or quadrinucleate). Staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate or tricolporate, heterocolpate (with pseudocolpi alternating with apertures), shed as monads, usually bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, echinate or psilate.

Gynoecium Pistil composed of two to five (to eight) connate carpels; carpels alternisepalous or odd carpel abaxial (sometimes with nectary at apex). Ovary usually inferior (rarely semi-inferior), unilocular (sometimes with nectary at apex). Style single, simple. Stigma usually punctate (sometimes capitate), non-papillate, Wet type. Pistillodia?

Ovules Placentation axile to apical. Ovules (one or) two to seven (to 20) per ovary, anatropous, pendulous, bitegmic, crassinucellar. Funicle long. Micropyle usually bistomal, Z-shaped (zig-zag; in at least one species of Guiera endostomal). Outer integument three to five cell layers thick. Inner integument two or three cell layers thick. Funicle long. Funicular obturator usually present. Hypostase often present. Parietal tissue eight to ten cell layers thick. Nucellar cap six to eight cell layers thick. Megagametophyte usually monosporous, Polygonum type (rarely tetrasporous, 16-nucleate, Penaea type). Synergids often with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis asterad. Vivipary occurring in mangrove species.

Fruit Usually a single-seeded, often two- to five-winged nutlike fruit (sometimes drupaceous, rarely dehiscent), often flattened.

Seeds Aril absent. Seed large. Exotesta? Endotesta tracheidal or sclerotic, without crystals? Exotegmen often fibrous. Endotegmen? Perisperm not developed. Endosperm absent. Embryo large, well differentiated, often with chlorophyll. Cotyledons one (by fusion), two or three (in some species of Terminalia up to five), usually convolute (sometimes conduplicate or plicate, sometimes hemispherical or spirally twisted). Germination phanerocotylar or cryptocotylar.

Cytology x = 7, 11–13 – Polyploidy occurring.

DNA

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavone-C-glycosides, 5-deoxyflavonoids, flavonoid sulphates, cyanidin, ellagic acid, galloyl- and ellagitannins, condensed tannins, alkaloids, triterpene saponins, cyanogenic compounds, and polyacetate derived arthroquinones present. Raffinose and stachyose present in phloem exudates.

Use Ornamental plants, timber, dyeing substances, tanning, fruits (Terminalia catappa), medicinal plants.

Systematics Combretaceae may be sister-group to the remaining Myrtales, although with fairly low support. Here, they are part of a basal trichotomy also including [Lythraceae+Onagraceae] and a clade comprising the remainder of Myrtales.

Strephonema is sister to the remaining Combretaceae.

Strephonematoideae Engl. et Diels in Engler, Monogr. Afr. Pflanzen-Fam. 3: 2. Nov 1899

1/3–4. Strephonema (3–4; tropical West and Central Africa). – Imperforate tracheary xylem elements with bordered pits. Intraxylary or interxylary phloem absent. Hairs furcate, adpressed. Leaves seemingly alternate. Stomata paracytic. Pollen grains without pseudocolpi. Exine semitectate, reticulate. Ovary semi-inferior (fruit largely superior). Ovules two per ovary. Cotyledons hemispherical, large, conduplicate. n = ?

Combretoideae (Lindl.) Beilschm. in Flora 16(Beibl. 7): 97. 14 Jun 1833 [‘Combreteae’]

11/490–525. Petiole sometimes glanduliferous. Pedicel often absent. Ovary inferior. Megagametophyte sometimes tetrasporous, 16-nucleate. Fruit flattened and/or winged (sometimes drupaceous). Cotyledons flattened and variously folded.

Laguncularieae Engl. et Diels in Engler, Monogr. Afr. Pflanzen-Fam. 3: 3. Nov 1899

4/10. Laguncularia (1; L. racemosa; coasts in tropical West Africa and tropical America), Lumnitzera (2; coasts in tropical East Africa, Madagascar and the Seychelles to islands in the Pacific), Macropteranthes (5; Northern Territory, eastern Queensland), Dansiea (2; eastern Queensland). – Pantropical, often mangroves. Lamina sometimes glanduliferous, in Laguncularia with revolute ptyxis. Stomata often cyclocytic. Floral prophylls (bracteoles) adnate to ovary. Petals sometimes clawed. Cotyledons spirally folded (convolute). n = 13. – The mangrove tree genera Lumnitzera and Laguncularia are sister-groups in the analyses by Maurin & al. (2010). On the other hand, Dansiea and Macropteranthes were not included in their analyses.

Combreteae DC., Prodr. 3: 18. Mar (med.) 1828

7/480–515. Terminalia (c 200; tropical regions on both hemispheres), Anogeissus (8; tropical Africa, southern Arabian Peninsula, tropical southern and Southeast Asia; in Terminalia?), Buchenavia (c 25; tropical America; in Terminalia?), Conocarpus (2; coasts in tropical Africa and tropical America); Getonia (1; G. floribunda; tropical Asia), Guiera (1; G. senegalensis; northern tropical Africa), Combretum (240–275; tropical regions on both hemispheres). – Pantropical. Usually trees (sometimes lianas). Vessel elements of two very different diametres, uniseriate wood rays and special radial vessels with perforations in tangential cell walls present in Combretum. Intraxylary phloem present. Mucilage ducts present in Terminalia. Stomata anomocytic. Petals sometimes small or absent. Micropyle in Guiera endostomal. Integuments sometimes multiplicative. Archespore sometimes multicellular. – Combretum is sister to the clade [Getonia+Guiera] (Maurin & al. 2010).

Cladogram of Combretaceae based on DNA sequence data (Maurin & al. 2010).

CRYPTERONIACEAE A. DC.

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A. de Candolle in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 16(2): 677. Jul 1868, nom. cons.

Henslowiaceae Lindl., Key Bot.: 57. 15-30 Sep 1835 [’Hensloviaceae’]; Henslowiales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 14. Sep-Oct 1835 [‘Henslowiaceae’]

Genera/species 3/10–13

Distribution Sri Lanka, northeastern India, southwestern China, the Andaman Islands, Indochina, Malesia and New Guinea.

Fossils Uncertain. Fossil pollen grains attributed to Dactylocladus have been found in Miocene layers on Borneo.

Habit Bisexual or dioecious (sometimes polygamodioecious), evergreen trees or shrubs. Young stems and branches often quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio usually deeply seated (in Dactylocladus subepidermal). Cortical vascular strands present or absent. Primary vascular tissue bicollateral. Vessel elements diffuse, with simple perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal aliform, confluent, reticulate, or banded (apotracheal parenchyma diffuse, diffuse-in-aggregates), rare or absent. Intraxylary phloem present. Sieve tube plastids Ss type? Nodes 3:3, trilacunar with trifid/split lateral vascular strands, or 1:3, unilacunar with three traces (with girdling vascular bundles). Sclereids often present. Crystals absent. Calciumoxalate raphides absent.

Trichomes Hairs usually absent (sometimes unicellular, simple).

Leaves Opposite, simple, entire, often coriaceous, with ? ptyxis. Stipules small lateral (Crypteronia), or absent; leaf sheath absent. Petiole vascular bundle transection arcuate or annular; petiole with wing bundles, sometimes with medullary leaf trace. Venation usually pinnate (rarely palmate), brochidodromous, with one continuous vein near leaf margin. Stomata usually paracytic (in Dactylocladus laterocytic to anemocytic). Cuticular wax crystalloids? Mesophyll usually with sclerenchymatous idioblasts (absent in Axinandra) and calciumoxalate styloids (and druses?). Leaf margin entire.

Inflorescence Terminal or axillary, panicle with long racemose or spicate branches.

Flowers Actinomorphic, small. Hypanthium tubular or cupular. Epigyny or half epigyny. Sepals four or five (or six), very small, with valvate aestivation, persistent or caducous, free or connate at base. Petals four or five (or six), very small, usually free, modified to caducous calyptra (Axinandra) or individually abscised (Crypteronia, Dactylocladus), or absent. Nectary? Disc absent.

Androecium Stamens four or five (or six; in Axinandra 5+5), haplostemonous or diplostemonous (Axinandra). Filaments inflexed in bud, free from each other and usually from tepals (in Axinandra epitepalous). Anthers basifixed, non-versatile, tetrasporangiate (microsporangia lateral to terminal), introrse to latrorse, longicidal (dehiscing by longitudinal slits); connective often distally (abaxially) expanded; endothecium ephemeral. Tapetum secretory. Staminodia absent?

Pollen grains Microsporogenesis simultaneous? Pollen grains tri- or tetracolporate, heterocolpate (in Axinandra, Dactylocladus with pseudocolpi alternating with apertures, in Crypteronia disyncolporate), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, rugulate or smooth.

Gynoecium Pistil composed of two to five (or six) connate carpels. Ovary inferior or semi-inferior, usually unilocular (septa sometimes fused at base). Style single, simple, usually filiform, often persistent. Stigma usually capitate (sometimes punctate), Wet type? Pistillodium absent?

Ovules Placentation parietal or basal-parietal. Ovules one to three (basal) to numerous (parietal) per carpel (six to 15 per ovary), anatropous, horizontal or ascending, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Transseptal vascular bundles sometimes present. Megasporangial tissue early degenerating. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule (with two to five valves, usually adnate to style), sometimes flattened.

Seeds Aril absent. Testa often winged. Exotestal cells tanniniferous. Endotesta multiplicative; endotestal cells crystalliferous. Exotegmic and endotegmic cells tanniniferous. Remaining layers more or less persistent. Perisperm not developed. Endosperm absent. Embryo straight, chlorophyll? Cotyledons two. Germination?

Cytology n = 8

DNA

Phytochemistry Virtually unknown. Aluminium accumulated.

Use Timber.

Systematics Dactylocladus (1; D. stenostachys; Borneo), Crypteronia (4–7; Assam, the Benghal, southwestern China, Indochina, the Andaman Islands, Malesia to New Guinea), Axinandra (5; Sri Lanka, West Malesia).

Crypteroniaceae are sister-group to the clade [Alzatea+[Rhynchocalyx+[Olinia+Penaeaceae]]].

Conti & al. (2002) recovered the topology [Dactylocladus+[Crypteronia+Axinandra]] for Crypteroniaceae.

LYTHRACEAE J. St.-Hil.

( Back to Myrtales )

Saint-Hilaire, Expos. Fam. Nat. 2: 175. Feb-Apr 1805 [‘Lythrariae’], nom. cons.

Salicariaceae Juss., Gen. Pl.: 330. 4 Aug 1789 [‘Salicariae’], nom. illeg.; Punicaceae Bercht. et J. Presl, Přir. Rostlin 2: 378. 1825 [‘Puniceae’], nom. cons.; Trapaceae Dumort., Anal. Fam. Plant.: 36, 39. 1829, nom. cons.; Ammanniaceae Horan., Prim. Lin. Syst. Nat.: 86. 2 Nov 1834 [‘Ammanniaceae (Lythrariae)’]; Trapales J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 549. 1846 [‘Trapaceae’]; Lagerstroemiaceae J. Agardh, Theoria Syst. Plant.: 338. Apr-Sep 1858 [‘Lagerstroemieae’]; Lawsoniaceae J. Agardh, Theoria Syst. Plant.: 338. Apr-Sep 1858 [‘Lawsonieae’]; Blattiaceae Engl., Syllabus: 146. Apr 1892; Sonneratiaceae Engl. in Engler et Prantl, Nat. Pflanzenfam. Nachtr. 1: 261. Oct 1897, nom. cons.; Duabangaceae Takht., Florist. Reg. World: 332. 27 Apr 1986

Genera/species 27/535–550

Distribution Cosmopolitan except polar areas, with their highest diversity in America.

Fossils Pollen grains assigned to Lythrum elkensis have been found in Lower Campanian strata in Wyoming. Seeds of Alatospermum, Microdiptera, Mneme and the extant Decodon are frequent in Cenozoic layers in North America and Europe. The palynogenus Florschuetzia from Cenozoic strata in Europe and Asia may be a predecessor of Sonneratia (and perhaps of Trapa). Lythrum and Trapa is known at least from the mid-Cenozoic onwards and the fossil genus Trapago from the Maastrichtian may possibly be attributed to Lythraceae.

Habit Usually bisexual (in Capuronia dioecious; in Sonneratia and Duabanga sometimes monoecious?), evergreen trees or shrubs, perennial or annual herbs (rarely lianas). Many species are hygrophytes, some representatives are aquatic (Sonneratia comprises mangrove trees with pneumatophores, producing vertical anchor roots, and horizontal nutrition roots; Trapa is aquatic). Bark often exfoliating. Young stems and branches usually quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio usually deeply (in Sonneratia and Duabanga superficially) seated. Polyderm present. Primary vascular tissue bicollateral. Secondary lateral growth normal or absent. Vessel elements usually with simple (in Sonneratia sometimes also scalariform and/or reticulate) perforation plates; lateral pits alternate or opposite, simple and/or bordered pits. Vestured pits present. Imperforate tracheary xylem elements usually fibre tracheids (in, e.g., Punica and Sonneratia) libriform fibres with simple or bordered pits (sometimes with crystals), septate or non-septate. Wood rays usually uniseriate (rarely multiseriate), usually heterocellular (rarely homocellular). Axial parenchyma apotracheal diffuse, or paratracheal scanty (sometimes vasicentric, aliform, confluent, or banded, or absent, i.a., in Punica and Sonneratia). Tyloses sometimes frequent. Intraxylary phloem present. Sieve tube plastids Ss type. Nodes usually 1:1 (or 1:3), unilacunar with one leaf trace (sometimes trilacunar). Sonneratia has branched sclereids in pneumatophores. Punica has secretory cells in cortex and medulla. Calciumoxalate as styloids, crystal sand, or prismatic or rhomboidal crystals.

Trichomes Hairs uni-, bi-, or multicellular, uniseriate or multiseriate, simple or branched (sometimes furcate, dendritic or stellate), or absent; multicellular glandular hairs present in some genera.

Leaves Usually opposite (sometimes verticillate or alternate, spiral; in Trapa with floating leaves in rosette and finely laciniate foliaceous submersed adventitious roots), simple, entire, often coriaceous, with conduplicate to flat ptyxis. Axillary glands usually present. Stipules small, early caducous, or absent; leaf sheath absent. Petiole in Trapa inflated, aerenchymatous. Petiole vascular bundle transection arcuate. Venation pinnate, usually brochidodromous. Stomata usually anomocytic (sometimes anisocytic, rarely paracytic; in Trapa adaxial). Cuticular wax crystalloids usually as platelets. Lamina sometimes gland-dotted, in several genera with hydathodes, salt-excreting glands, or nectaries. Epidermis often with large mucilaginous idioblasts. Mesophyll usually without sclerenchymatous idioblasts (present in Sonneratia and Duabanga), often with calciumoxalate as druses or solitary prismatic crystals (in Punica with large prismatic crystals). Sclereids present in Duabanga and asterosclereids in Sonneratia. Leaf margin usually entire (sometimes serrate or dentate). Extrafloral nectaries rarely (e.g. in Lafoensia) present on lamina.

Inflorescence Terminal or axillary, cyme, thyrse, corymb, umbel- or head-like, spike or raceme (flowers sometimes solitary axillary).

Flowers Actinomorphic or zygomorphic. Hypanthium cupular or tubular (sometimes with spurs), often distinctly ribbed, or absent. Usually hypogyny or half epigyny (rarely epigyny). Androphore present in some genera. Sepals (three or) four to eight (to 16), with valvate aestivation, often with alternate appendices (epicalyx), often persistent and finally coriaceous, free. Petals (three or) four to eight (to 16; in Decodon usually five), with plicate or imbricate aestivation, often wrinkled in bud, often clawed, inserted at adaxial margin of hypanthium, caducous, free (absent in some species). Nectariferous disc annular or unilateral intrastaminal at ovary base, or absent (in Cuphea often unilateral gland). Diheterostyly and triheterostyly occurring in some genera (e.g. Lythrum).

Androecium Stamens one to more than 100 (in Decodon usually 5+5; in Cuphea 11; in Trapa four), in one, two or three whorls, sometimes as many as sepals, antesepalous or antepetalous, usually obdiplostemonous (sometimes diplostemonous or haplostemonous), centripetally or centrifugally developing. Filaments free from each other and from tepals, inserted immediately below petals to near ovary, with unequal lengths (when diplostemonous then antesepalous filaments longer, inserted near or above base of perianth tube). Anthers sometimes inflexed in bud, usually dorsifixed (rarely basifixed), usually versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to sexanucleate (in Trapa multinucleate) cells. Staminodia usually absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate or tricolporate (in Duabanga triporate; in Trapa trichotomocolpate; in Sonneratia?), usually heterocolpate (with three or six pseudocolpi alternating with apertures), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, psilate, scabrate or finely verrucate or striate (in Trapa with three distinct ridges running transversely across colpi and fusing at poles).

Gynoecium Pistil composed of two (in, e.g., Trapa, transversely orientated) to six (to c. 15) connate carpels; when carpels as many as sepals, then alternisepalous or antesepalous; when carpels two, then often transverse or median; when carpels three, then median carpel adaxial. Ovary superior to semi-inferior (rarely inferior), usually bilocular to quadrilocular (rarely unilocular or multilocular), with septa incomplete at apex or reduced to thin threads. Style single, simple, thin (in Trapa hollow). Stigma punctate or capitate, papillate, Dry or Wet type (Lagerstroemia, Punica). Pistillodium?

Ovules Placentation usually axile (almost free central at maturation; rarely parietal; in Trapa apical-axile; carpels in Punica granatum in two or three superposed layers, basal carpels with axile and remaining carpels with intrusively parietal placentation). Ovules usually two to c. 50 per carpel, anatropous, ascending or horizontal (in Trapa one, pendulous), bitegmic, crassinucellar. Micropyle usually bistomal (in Punica exostomal?; micropyle in Trapa formed by long and massive nucellar beak). Outer integument two to seven (to nine) cell layers thick. Inner integument two or three cell layers thick. Hypostase usually present. Parietal tissue approx. seven cell layers thick. Megasporocytes several (archespore multicellular). Megagametophyte monosporous, Polygonum type. Antipodal cells not formed in Trapa. Endosperm development ab initio nuclear (endosperm not developing in Trapa, suspensor haustorium formed). Endosperm haustoria? Embryogenesis usually onagrad (in Trapa solanad).

Fruit Usually a loculicidal, septicidal or irregularly dehiscing capsule (sometimes a pyxidium; rarely, e.g. in Punica, a berry; in Trapa a one-seeded nutlike fruit with persistent sepals modified into hard horn-like processes; fruit in Cuphea dehiscing by placenta penetrating pericarp and hypanthium), often more or less enclosed by persistent hypanthium and/or sepals. Placenta sometimes expanded in fruit.

Seeds Aril absent. Seed usually flat. Operculum often? present. Testa usually multiplicative, multi-layered, sometimes winged, usually with epidermal internal finger-shaped initially inverted invaginations of plasmalemma developing into mucilaginous hairs when wet. Exotesta of various shape (seeds in Punica embedded in pulpy tissue formed by enlarged epidermal exotestal cells). Endotestal cells often sclerotic, often tracheidal (sometimes crystalliferous). Exotegmen and endotegmen sometimes consisting of interwoven fibres (in Punica undistinguished). Perisperm not developed. Endosperm usually absent (in Trapa present, starchy). Spiral suspensor present in Trapa. Embryo usually straight, well differentiated, usually without (in Sonneratia and Duabanga with) chlorophyll. Cotyledons two, oily, usually flat (in Lagerstroemia and Punica plicate; in Trapa very unequal in size). Radicula absent in Trapa. Germination phanerocotylar (in Trapa also cryptocotylar; large starchy cotyledon persisting inside pericarp, whereas small cotyledon, plumule and radicula penetrate apical pore arising when style detached).

Cytology n = (5–)8(–15, 24, 28, 32), n = 12 (Duabanga, Sonneratia); n = 20, 24 (Trapa); n = 6–c. 86 (Cuphea); x = 8 – Polyploidy and aneuploidy frequently occurring.

DNA

Phytochemistry Flavonols (kaempferol, quercetin), ellagic acid, tannins (e.g. cuphiin, ellagitannins), quinolizidine alkaloids, and polyacetate derived arthroquinones present. Proanthocyanidins, saponins and cyanogenic compounds not found. Raffinose and stachyose present in phloem exudates. Dissolvable oxalate accumulated.

Use Ornamental plants, fruits (Punica granatum, Trapa), vegetables (Trapa), dyeing substances (Lawsonia inermis), medicinal plants, timber.

Systematics Lythraceae are sister-group to Onagraceae.

Lythroideae are sister to the clade [Lagerstroemioideae+Punicoideae] (Graham & al. 2011).

Lythroideae Juss. ex Arn., Botany: 108. 9 Mar 1832 [‘Lythrarieae’]

4/c 85. Lythrum (36; nearly cosmopolitan), Decodon (1; D. verticillatus; eastern United States); Rotala (c 45; temperate to tropical regions on both hemispheres), Heimia (2–3; southern Texas, southern New Mexico, Mexico, Central America, the West Indies, South America to northern Argentina). – Subcosmopolitan.

[Lagerstroemioideae+Punicoideae]

Testa with inverted mucilaginous hairs.

Lagerstroemioideae Beilschm. in Flora 16(Beibl. 7): 99, 108. 14 Jun 1833 [’Lagerstroemidae’]

9/100–105. Lagerstroemia (50–55; tropical Asia, northern and eastern Australia), Duabanga (2; tropical Asia), Sonneratia (6; coasts of the Indian and Pacific Oceans), Trapa (1; T. natans; warm-temperate regions in Europe, Africa and Asia); Ammannia (c 25; nearly cosmopolitan), Crenea (2; Trinidad, tropical South America), Ginoria (14; Mexico, the West Indies), Tetrataxis (1; T. salicifolia; Mauritius), Lawsonia (1; L. inermis; North Africa, tropical regions in the Old World). – Nearly cosmopolitan. Sonneratia consists of mangrove trees. – Lawsonia, Ammannia, Crenea, Ginoria and Tetrataxis may form a clade.

Punicoideae (Eichl.) Luerss., Handb. Syst. Bot. 2: 822. Oct 1881 [‘Puniceae’]

14/350–360. Woodfordia (2; W. uniflora: northeastern Africa, southern Arabian Peninsula; W. fruticosa: Madagascar, tropical Asia eastwards to southern China and Timor), Koehneria (1; K. madagascariensis; southern Madagascar), Cuphea (c 260; America), Pleurophora (7–10; South America); Lafoensia (5–6; tropical America), Pemphis (1; P. acidula; tropical regions in the Old World), Punica (2; P. protopunica: Socotra; P. granatum: southwestern Asia), Adenaria (1; A. floribunda; southern Mexico to Panamá, eastern South America to northern Argentina), Pehria (1; P. compacta; Colombia, Venezuela), Capuronia (1; C. madagascariensis; Madagascar), Galpinia (1; G. transvaalica; eastern southern Africa northwards to Zimbabwe and Mozambique), Diplusodon (65–70; Brazil), Lourtella (1; L. resinosa; Peru), Physocalymma (1; P. scaberrimum; tropical South America). – Pantropical, southeastern Africa, southwestern Asia, warm-temperate and subtropical America. – Koehneria may be sister to Woodfordia. Cuphea and Pleurophora are sister-groups. Adenaria may be sister-group to Pehria. Capuronia and Galpinia are sister-groups.

Maximum likelihood tree of Lythraceae based on DNA sequence data (Graham & al. 2011).

MELASTOMATACEAE Juss.

( Back to Myrtales )

de Jussieu, Gen. Plant.: 328. 4 Aug 1789 [’Melastomae’], nom. cons.

Melastomatales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 233. Jan-Apr 1820 [‘Melastomeae’]; Blakeaceae Reichb. ex Barnhart in Bull. Torrey Bot. Club 22: 19. 15 Jan 1895; Miconiaceae Mart., Consp. Regn. Veg.: 64. Sep-Oct 1835 [’Miconieae’]; Rhexiaceae Dumort., Comment. Bot.: 59. Nov-Dec 1822 [‘Rhexideae’]

Genera/species c 154/4.400–-4.625

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

Fossils Uncertain. Fossil flowers, possibly attributable to Melastomataceae, have been found in Turonian and younger layers (Crepet 2008).

Habit Usually bisexual (rarely androdioecious), evergreen trees or shrubs, perennial herbs (sometimes aquatic) or lianas. Many species are epiphytic. Young stems and branches often quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Cortical and medullary vascular bundles present or absent. Primary vascular tissue bicollateral or centrifugal. Secondary lateral growth normal, anomalous (via cylindrical cambium) or absent. Vessel elements with usually simple (sometimes reticulate) perforation plates; lateral pits alternate, usually simple. Vestured pits present. Imperforate tracheary xylem elements usually libriform fibres (in, e.g., Pternandra fibre tracheids) with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, and/or paratracheal scanty, reticulate, scalariform, vasicentric or banded. Interxylary phloem usually absent (present in Pternandra). Sieve tube plastids Ss type. Endodermis sometimes prominent (rarely with corky cell walls). Nodes usually 1:1, unilacunar with one leaf trace (sometimes 1:3, unilacunar with trifid/split lateral vascular strands). Cortex usually without cristarque cells (present in Osbeckieae). Heartwood sometimes with gum-like substances. Prismatic calciumoxalate crystals, druses and/or crystal sand present or absent. Calciumoxalate raphides absent. Root tips with anthocyanins.

Trichomes Hairs usually large, multicellular, uniseriate or multiseriate, simple, stellate, peltate (especially in Astronieae) or often lepidote or vesicular, often dendritic; multicellular glandular hairs (sometimes lepidote) present (Melastomateae with characteristic ’tibouchinoid’ hairs).

Leaves Usually opposite (rarely verticillate), simple, entire, with conduplicate or supervolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate, annular etc. Venation palmate-parallelodromous, acrodromous (rarely pinnate); primary veins three to nine (to 19), arising arcuately at or near leaf base and confluent at leaf tip; tertiary veins usually prominent, scalariform, running at right angle against midvein. Stomata usually anomocytic (sometimes anisocytic, paracytic, polycytic, diacytic, cyclocytic or anomocyclocytic). Cuticular wax crystalloids? Domatia usually absent (in, e.g., Blakeeae and Miconieae often acarodomatia or myrmecodomatia as pits or pockets). Epidermis with or without mucilaginous idioblasts. Mesophyll usually with sclerenchymatous idioblasts containing sclereids of various types and with calciumoxalate as raphides, styloids, druses and crystal sand. Secretory cavities absent. Leaf margin usually entire (sometimes serrate).

Inflorescence Terminal or axillary, panicle, thyrse or other types of cymose inflorescences (flowers rarely solitary terminal). Bracts and floral prophylls (bracteoles) often showy.

Flowers Actinomorphic or zygomorphic (especially by androecium). Hypanthium tubular or campanulate. Epigyny or half epigyny. Sepals four or five (to seven), with imbricate, valvate, contorted or open aestivation, on margin of hypanthium, often early caducous, usually free (sometimes connate into caducous calyptra). Petals four or five (to seven), with destrorsely-contorted aestivation, on margin of hypanthium, free. Nectary usually absent (nectar in some genera secreted from hypanthium, petal tips, filaments, anther connectives, ovary apex, or stigma). Disc absent.

Androecium Stamens usually 3+3, 4+4, or 5+5 (rarely three or up to more than 100), usually diplostemonous (rarely obdiplostemonous or haplostemonous); inserted on one side of flower. Filaments often twisted, often geniculate at connective base, free from each other and from tepals. Anthers sometimes inflexed in bud, inverted during development, basifixed, non-versatile, tetrasporangiate, introrse, usually poricidal (dehiscing by one, two or four apical pores, developing in epidermis-free part of microsporangia by desiccation of exposed mesophyll; fibre thickenings absent in endothecium; in Pternandra and Astronieae introrse?, longicidal, with short or long longitudinal slits); connective enlarged in characteristic way, often basally-ventrally prolonged or with horn- or spur-like dorsal appendages, without glands. Tapetum secretory, with ?-nucleate cells. Staminodia usually absent (sometimes four or five, alternating with fertile stamens).

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colpor(oid)ate or 3(–6)-colpate, heterocolpate (usually with three pseudocolpi or intercolpate depressions alternating with apertures), usually shed as monads (rarely as tetrads or polyads), bicellular at dispersal. Exine tectate, with columellate infratectum, perforate or punctate, striate, rugulate, rugulate-verrucate or smooth.

Gynoecium Pistil composed of (three or) four or five (to 14) connate usually antesepalous carpels. Ovary inferior or semi-inferior, (unilocular or) quadri- or quinquelocular (to 14-locular), usually with antesepalous (in Pternandra and Rhexia antepetalous) locules, often with interspace between ovary wall and hypanthium. Style single, simple, sometimes hollow. Stigma punctate to capitate, papillate, Wet type. Pistillodium?

Ovules Placentation usually axile (in Astronieae basal; in Pternandra parietal). Ovules usually numerous (sometimes one or few) per carpel, usually anatropous (in Rhexia orthotropous), bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument approx. two cell layers thick. Inner integument two or three cell layers thick. Hypostase present. Parietal tissue four to six cell layers thick. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad or solanad.

Fruit A loculicidal capsule, with walls often degenerating during maturation and leaving vertical vascular bundles (capsule sometimes dehiscing along epigynous part or irregularly), or a berry.

Seeds Aril absent. Small seeds with operculum formed by hilum. Exotesta palisade to cuboid and lignified. Mesotesta sclerotic. Endotesta? Tegmen crushed, without fibrous exotegmen. Perisperm not developed? Endosperm absent. Embryo small, well differentiated, (with? or) without chlorophyll. Cotyledons two, small, often unequal in size. Radicula curved, inserted in testal pouch. Germination phanerocotylar.

Cytology n = (8–)9–17 (23, 31) – Polyploidy frequently occurring.

DNA

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, delphinidin, acylated anthocyanins, ellagic acid, tannins, and phenylalanine-derived cyanogenic compounds present. Saponins not found. Aluminium accumulated at least in Pternandra, Astronieae, Merianieae, and Miconieae.

Use Ornamental plants, fruits, timber, dyeing substances.

Systematics Melastomataceae are probably sister to Memecylaceae.

Pternandra (Kibessioideae) is sister to the remaining Melastomataceae.

Kibessioideae Naudin in Ann. Sci. Nat. Bot., sér. 3, 12: 201. Oct 1849 [‘Kibessieae’]

1/1–15. Pternandra (1–15; Indochina, Hainan, Malesia and eastwards to northern Queensland). – Phellogen superficial. Internal phloem present. Hairs uniseriate. Interpetiolar stipular flange present below leaves. Petiole vascular bundle transection arcuate. Stomata anomocyclocytic. Flowers tetramerous. Endothecium present only in internal wall of inner microsporangium. Anther connectives with apical oil-glands. Carpels antepetalous. Ovary divided by secondary septa. Placentation parietal. Capsule carnose. Aluminium accumulated.

Melastomatoideae Ser. ex DC., Prodr. 3: 100. Mar (med.) 1828 [‘Melastomeae’]

c 153/4.400–4.610. Internal phloem absent. Petiole vascular bundle transection of various types. Leaf veins without fibrous envelope. Inflorescence often terminal. Sepals sometimes connate, calyptrate. Anthers poricidal (dehiscing by apical pores). Endothecium ephemeral or absent. Carpels usually antesepalous, developing before anthers.

Astronieae Triana in Bull. Cong. Int. Bot. Hort. Amsterdam 1865: 457. 1866

3–4/c 150. Astrocalyx (1; A. calycina; the Philippines), Astronia (59; tropical Asia and eastwards to islands in the Pacific), Astronidium (67; Borneo and the Philippines to New Guinea, Fiji, Micronesia to the Society Islands; incl. Beccarianthus?), Beccarianthus (>22; Borneo to New Guinea; in Astronidium?). – Tropical Asia and eastwards to the Society Islands. Phellogen superficial. Petiole vascular bundles complex, open. Stomata usually anomocytic. Hairs peltate-lepidote. Anthers sometimes longicidal (dehiscing by slits). Carpels antepetalous. Placentation basal to basal-axile. Aluminium accumulated. – Astronieae are sister-group to the remaining Melastomatoideae.

[[Macrocentrum+[Merianieae+[Miconieae+Henrietteeae]]]+[Bertolonieae+[[Blakeeae+Sonerileae]+[Microlicieae+[Rhexieae+Melastomateae]]]]]

Highest diversity in tropical and subtropical South America. Trees or herbs (sometimes epiphytes, rarely lianas). Phellogen sometimes superficial. Hair types diverse (including short-stalked glands). Petiole vascular bundle transection arcuate or complex. Secondary veins three or five from (near) base; tertiary veins perpendicular to mid-vein. Stomata polycytic, cyclocytic, etc. Flowers (3–)4–5(–10)-merous. Calyx sometimes with alternate lobes. Stamens sometimes as many as sepals, antesepalous or antepetalous, numerous. Anthers poricidal. Connective often with basal appendage, sometimes nectariferous. Carpels two to 15. Placentation axile. Ovules sometimes one to few per carpel. Style sometimes hollow. Capsule sometimes irregularly dehiscent (sometimes a berry). Aluminium accumulated in Merianieae and Miconieae.

[Macrocentrum+[Merianieae+[Miconieae+Henrietteeae]]]

Macrocentrum clade

1/15–20. Macrocentrum (15–20; Central America and northern tropical South America).

[Merianieae+[Miconieae+Henrietteeae]]

Merianieae Triana in Bull. Cong. Int. Bot. Hort. Amsterdam 1865: 457. 1866

c 16/230–240. Adelobotrys (25–30; tropical America), Axinaea (25–30; tropical America; in Meriania?), Behuria (14; southern Brazil), Benevidesia (2; southeastern Brazil), Bisglaziovia (1; B. behurioides; Brazil), Centronia (15; Central America, western tropical South America, Guianas), Dolichoura (1; D. spiritusanctensis; Brazil), Graffenrieda (44; tropical America), Huberia (c 14; Ecuador, Peru, southeastern Brazil), Meriania (74; tropical America), Merianthera (3; Brazil), Neblinanthera (1; N. cumbrensis; Venezuela), Ochthephilus (1; O. repentinus; Guyana), Phainantha (4; tropical South America), Tessmannianthus (6–7; tropical America). – Tropical America.

[Miconieae+Henrietteeae]?

Miconieae DC., Prodr. 3: 152. Mar (med.) 1828

c 24/1.620–1.670. Allomaieta (8; Colombia), Alloneuron (7; Colombia, Peru), Quipuanthus (1; Q. epipetricus; the Andes in Ecuador and Peru), Wurdastom (8; tropical South America); Anaectocalyx (3; Venezuela), Boerlagea (1; B. grandiflora; Borneo), Calycogonium (c 35; the West Indies), Catocoryne (1; C. linnaeoides; Peru), Charianthus (6; the Lesser Antilles), ’Clidemia’ (115–120; tropical America; non-monophyletic), Conostegia (43; tropical America, with their highest diversity in Central America), Creochiton (9; Central and East Malesia), Eriocnema (1; E. fulva; Brazil), Physeterostemon (3; Bahia in Brazil), Killipia (5; Colombia), ’Leandra’ (175–200; tropical America, with their highest diversity in Brazil; non-monophyletic?), Maieta (3; tropical America), Mecranium (24; the West Indies), ’Miconia’ (c 1.000; tropical America; non-monophyletic), Ossaea (80–90; tropical America; polyphyletic?), Pachyanthus (16; Cuba, Hispaniola, Colombia), Pleiochiton (7; southern Brazil), Pseudodissochaeta (6; northern India to Hainan), Tetrazygia (20–25; Florida, the West Indies), ’Tococa’ (47–54; tropical America; non-monophyletic). – Tropical Asia, tropical America. – Allomaieta (including Cyphostyla), Alloneuron and Wurdastom should possibly be transferred to Cyphostyleae (Michelangeli & al. 2011). They have antesepalous stamens, inferior ovary, and capsule-like fruit. Yet, will this render Miconieae paraphyletic?

Henrietteeae Penneys, Michelang., Judd et Almeda in Syst. Bot. 35(4): 795. 6 Dec 2010

3/77. Bellucia (8; tropical America), Henriettea (67; tropical South America), Kirkbridea (2; Colombia). – Tropical America.

[Bertolonieae+[[Blakeeae+Sonerileae]+[Microlicieae+[Rhexieae+Melastomateae]]]]

Bertolonieae Triana in Bull. Cong. Int. Bot. Hort. Amsterdam 1865: 457. 1866 [‘Bertoloniceae’]

3/47–49. Bertolonia (17; Venezuela, southeastern Brazil), Monolena (8–10; tropical South America), Triolena (22; tropical America). – Tropical America.

[[Blakeeae+Sonerileae]+[Microlicieae+[Rhexieae+Melastomateae]]]

[Blakeeae+Sonerileae]

Blakeeae Benth. et Hook. f., Gen. Plant. 1: 727, 735. Sep 1867 (in Sonerileae?)

3/c 105. Blakea (c 100; tropical America), Chalybea (1; C. peruviana; tropical America), Huilaea (4; Colombia). – Tropical America.

Sonerileae Triana in Bull. Cong. Int. Bot. Hort. Amsterdam 1865: 457. 1866

c 43/1.040–1.070. Amphiblemma (13; tropical West and Central Africa), Anerincleistus (30; India and southern China to the Philippines), Aschistanthera (1; A. cristanthera; Vietnam), Barthea (1; China, Taiwan), Blastus (12; Assam to West Malesia), Boyania (1; B. ayangannae; Guianas), Bredia (c 30; East and Southeast Asia), Calvoa (19; tropical Africa), Catanthera (17; Sumatra, Borneo, New Guinea), Centradeniastrum (2; western tropical South America), Cincinnobotrys (7; tropical Africa), Cyphotheca (1; C. montana; Yunnan), Dicellandra (3; tropical West and Central Africa), Diplarpea (1; D. paleacea; Colombia), Diplectria (11; Southeast Asia, Malesia), Dissochaeta (22; tropical Asia), Driessenia (14–18; West and Central Malesia, with their largest diversity on Borneo), Feliciadamia (1; F. stenocarpa; French Guinea), Fordiophyton (14; southern China, Southeast Asia), ’Gravesia’ (c 110; tropical Africa, Madagascar; non-monophyletic), Kendrickia (1; K. walkeri; southern India, Sri Lanka), Kerriothyrsus (1; K. tetrandrus; Laos), Macrolenes (15; Thailand, Malesia), Maguireanthus (1; M. ayangannae; Guyana), ’Medinilla’ (375–400; tropical Africa, Madagascar, tropical Asia to southern China, Taiwan and New Guinea, northeastern Queensland, Fiji, Samoa; non-monophyletic), Neodriessenia (6; Borneo), Ochthocharis (7; tropical Africa, tropical Asia), Opisthocentra (1; O. clidemioides; northern Brazil), Oxyspora (c 25; tropical Asia, southern China), Pachycentria (8; Burma, Malesia), Phyllagathis (c 55; southern China, Southeast Asia, West Malesia), Plethiandra (9; West Malesia), Poikilogyne (21; Borneo, New Guinea), Poilannammia (4; Vietnam), Preussiella (2; tropical West and Central Africa), Salpinga (8; tropical South America), Sarcopyramis (1–3; tropical Asia), Scorpiothyrsus (6; Hainan, Southeast Asia), Sonerila (c 175; tropical Asia), Sporoxeia (6; Burma, Southeast Asia), Stanmarkia (2; Mexico, western Guatemala), Stussenia (1; S. membranifolia; Vietnam), Tateanthus (1; T. duidae; Venezuela), Tryssophyton (1; T. merumense; Guyana). – Pantropical. Aluminium accumulated in some species. – The position of Cambessedesia is not fully clarified.

[Microlicieae+[Rhexieae+Melastomateae]]

Connective with basal appendage (pedoconnective). – Michelangeli & al. (2013) found that Cambessedesia (21; eastern Brazil) is sister to a clade comprising Microlicieae, Rhexieae and Melastomateae. Furthermore, in their analyses Melastomeae were sister-group to [Microlicieae+Rhexieae].

Microlicieae Naudin in Ann. Sci. Nat. Bot., sér. 3, 12: 203. Oct 1849 [‘Microliciales’]

7/c 255. Rhynchanthera (15; Mexico to Bolivia, Paraguay and eastern Brazil); Trembleya (11; southern and eastern Brazil), Chaetostoma (12; Brazil), Lavoisiera (46; Brazil, with their largest diversity in eastern Brazil), Lithobium (1; L. cordatum; Brazil)?, Microlicia (c 170; tropical South America, with their highest diversity in southern and eastern Brazil), Stenodon (1; S. suberosus; southern Brazil). – Tropical America, with their highest diversity in the cerrado in southern and eastern Brazil.

[Rhexieae+Melastomateae]

Seeds cochleate.

Rhexieae DC., Prodr. 3: 114. Mar (med.) 1828

3/23–26. Arthrostemma (4–7; southern Mexico, Central America, the West Indies, South America to Bolivia), Pachyloma (6; Amazonia), Rhexia (13; eastern North America). – Tropical America.

Melastomateae Bartl., Ord. Nat. Plant.: 329. Sep 1830 [‘Melastomea’]

c 46/840–950. Acanthella (2; tropical South America), ‘Acisanthera’ (17; tropical America; polyphyletic), Siphanthera (16; northeastern South America), ‘Comolia’ (22; tropical South America; polyphyletic), Sandemania (1; S. hoehnei; Venezuela, Peru, Amazonian savannas in Brazil), Macairea (22; tropical South America), ‘Ernestia’ (16; tropical South America; polyphyletic), Nepsera (1; N. aquatica; tropical America), Appendicularia (1; A. thymifolia; Guianas), Marcetia (27; tropical South America), Fritzschia (1; F. erecta; southeastern Brazil), ‘Aciotis’ (13; tropical America; polyphyletic), ‘Pterolepis’ (14; tropical America, with their highest diversity in Brazil; paraphyletic; incl. Pterogastra?), Pterogastra (2; northern tropical South America; in Pterolepis?), Guyonia (2; tropical West and Central Africa), Tristemma (15; tropical Africa, Madagascar, the Mascarene Islands), Dichaetanthera (c 35; tropical Africa, Madagascar), Heterotis (6; tropical Africa), Antherotoma (2; tropical Africa, Madagascar; incl. Dissotis multiflora), Amphorocalyx (5; Madagascar), Dionycha (3; Madagascar), ‘Dissotis’ (c 120; tropical and southern Africa, Madagascar; polyphyletic), Osbeckia (c 50; Southeast Asia, Malesia to New Guinea, northern Australia), Melastoma (20–25; Southeast Asia, Malesia, northern and eastern Australia), Desmoscelis (2; Brazil), Heterocentron (28; southern Mexico, Central America), Centradenia (6; southern Mexico, Central America), Pilocosta (5; tropical America), Bucquetia (3; the Andes), Castratella (2; the Andes), Chaetolepis (10; the Andes), Monochaetum (c 45; Central America, the Andes), Brachyotum (58; the Andes; in Tibouchina?), Svitramia (6; southeastern Brazil; in Tibouchina?), Tibouchinopsis (2; northeastern Brazil; in Tibouchina?), ‘Tibouchina’ (240–350; Central America, the West Indies, South America; non-monophyletic). – unplaced: Cailliella (1; C. praerupticola; tropical West Africa), Comoliopsis (1; C. neblinae; Venezuela), Dinophora (2; tropical West and Central Africa), Dionychastrum (1; D. schliebenii; Uluguru Mountains in Tanzania), Loricalepis (1; L. duckei; Brazil), Mallophyton (1; M. chimantense; Venezuela), Melastomastrum (6; tropical Africa), Microlepis (4; southern Brazil), Nerophila (1; tropical West Africa; in Chaetolepis?), Poteranthera (2; tropical South America), Pseudosbeckia (1; P. swynnertonii; tropical East Africa), Schwackaea (1; S. cupheoides; southern Mexico, Central America). – Pantropical. Aluminium accumulated in some species. – Possibly, Melastomateae is sister to Rhexieae. The clade [Aciotis+Nepsera] may be sister-group to the remaining Melastomateae.

Unplaced Melastomataceae

Dalenia (3-6; Borneo), Vietsenia (4; Vietnam).

Cladogram (simplified) of Melastomataceae and Memecylaceae based on DNA sequence data (Clausing & al. 2000; Renner & al. 2001; Fritsch & al. 2004; Renner & al., unpubl.).

MEMECYLACEAE DC.

( Back to Myrtales )

de Candolle in D. F. L. von Schlechtendal in Linnaea 2: 505. Aug-Oct 1827 [‘Memecyleae’]

Memecylales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 65. Sep-Oct 1835 [‘Memecyleae’]; Mouriraceae Gardner in J. Bot. (Hooker) 2: 22. Feb 1840 [’Mouririaceae’]

Genera/species 6/280–290

Distribution Tropical Africa, Madagascar, Mauritius, Malesia, Central America, the West Indies and tropical South America.

Fossils Unknown.

Habit Usually bisexual (in Lijndenia androdioecious), evergreen trees or shrubs, or perennial herbs. Young stems and branches terete.

Vegetative anatomy Phellogen ab initio superficially or deeply seated? Primary vascular tissue often bicollateral. Secondary lateral growth normal or anomalous (via cylindrical cambium). Vessel elements with simple perforation plates; lateral pits alternate, usually (half-)bordered (rarely simple) pits. Vestured pits? Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually paratracheal scanty, reticulate, scalariform, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, or narrowly banded (sometimes apotracheal diffuse or diffuse-in-aggregates). Tyloses abundant. Intraxylary phloem foraminate, diffuse. Interxylary phloem present in secondary xylem. Sieve tube plastids Ss type. Nodes swollen, usually 1:1, unilacunar with one leaf trace (sometimes 1:3, unilacunar with three traces). Heartwood often with gum-like substances. Sclereids usually present. Calciumoxalate as acicular crystals and/or styloids usually present (sometimes crystal sand). Calciumoxalate raphides absent. Root tips without anthocyanins.

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

Leaves Opposite, simple, entire, with flat (Memecylon) or revolute (Mouriri) ptyxis. Stipules usually absent (present in seedlings); leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation usually palmate-parallelodromous (sometimes pinnate), acrodromous or (often seemingly brochidodromous), with arcuate main-veins and indistinct lateral veins; usually with fibrous envelope; in Memecylon and Mouriri with large sclereids at vein tips. Stomata paracytic or anomocytic. Cuticular wax crystalloids? Lamina sometimes pellucid-punctate. Mesophyll with or without sclerenchymatous idioblasts. Secretory cavities absent. Leaf margin entire.

Inflorescence Axillary, often fasciculate or umbel-like.

Flowers Actinomorphic or zygomorphic, small. Pedicel articulated. Hypanthium tubular or campanulate. Epigyny. Sepals (three or) four or five, with usually valvate (sometimes imbricate quincuncial or open) aestivation, inserted on margin of hypanthium, often caducous at beginning of anthesis, free. Petals (three or) four or five, with dextrorsely contorted aestivation, inserted on margin of hypanthium, free. Nectary absent. Disc absent.

Androecium Stamens usually 3+3, 4+4 or 5+5 (sometimes four or five). Filaments free, often bent in bud (in Votomita straight), free from tepals. Anthers dorsifixed, versatile, ab initio tetrasporangiate, as mature disporangiate, extrorse (inverted during development), poricidal (dehiscing by apical pores) or longicidal (dehiscing by longitudinal, sometimes short, slits, formed by irregular desiccation of endothecial cells); connective fleshy, usually with depressed elliptic terpenoid-secreting dorsal oil gland. Tapetum secretory, usually with uninucleate (not in Mouriri) cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tri- or hexacolpate and/or tri- or hexacolporate, heterocolpate (usually with three pseudocolpi or intercolpate depressions alternating with apertures), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, perforate, punctate, rugulate or smooth.

Gynoecium Pistil composed of (two to) four or five (to 14) connate carpels. Ovary inferior, unilocular to quinquelocular, with antepetalous locules. Style single, simple. Stigma punctate to capitate?, papillate?, Wet type. Pistillodium?

Ovules Placentation free central (when ovary unilocular) or axile-basal (when ovary multilocular). Ovules (one or) two to 18 (or more) per carpel, usually campylotropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument usually approx. two (in Mouriri four or five) cell layers thick. Inner integument approx. two cell layers thick. Parietal tissue approx. three cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A one- or several-seeded berry.

Seeds Aril absent. Operculum? Exotestal cells longitudinally elongated (in Memecylon few sclerotic hypodermal exotestal cells). Endotesta? Exotegmen fibrous. Endotegmen? Subhilum massively sclerotic. Perisperm developed, starchy? Endosperm absent. Embryo usually large, well differentiated, with chlorophyll. Cotyledons two, thick, straight or curved, lobate (in Memecylon crumpled). Radicula curved. Hypocotyl often elongating during seed development. Germination cryptocotylar or phanerocotylar.

Cytology n = 7

DNA

Phytochemistry Insufficiently known. Non-acylated anthocyanins and cyanogenic compounds present. Saponins not found. Aluminium accumulated.

Use Carpentries, dyeing substances.

Systematics Memecylon (c 150; tropical regions in the Old World), Warneckea (31–33; tropical Africa, Madagascar, Mauritius), Lijndenia (10; tropical regions in the Old World), Spathandra (6; tropical Africa, Madagascar), Mouriri (81; tropical America), Votomita (6–8; tropical America).

Memecylaceae are sister to Melastomataceae.

Pternandra in Melastomataceae was sister to [Mouriri+Memecylon], according to some ndhF analyses, a fact that should render Memecylaceae an ingroup in Melastomataceae.

Phylogeny (maximum-likelihood tree) of Memecylaceae based on DNA sequence data (Stone 2006).

MYRTACEAE Juss.

( Back to Myrtales )

de Jussieu, Gen. Plant.: 322. 4 Aug 1789 [’Myrti’], nom. cons.

Melaleucaceae Vest, Anleit. Stud. Bot.: 268, 286. 1818 [’Melaleucoideae’]; Baeckeaceae Bercht. et J. Presl, Přir. Rostlin: 233. Jan-Apr 1820 [‘Bekeae’]; Eugeniaceae Bercht. et J. Presl, Přir. Rostlin 2: 378. 1825; Leptospermaceae Bercht. et J. Presl, Přir. Rostlin 2: 378. 1825; Lythrales Link, Handbuch 2: 47. 4-11 Jul 1829 [‘Lythrariae’]; Chamelauciaceae DC. ex F. Rudolphi, Syst. Orb. Veg.: 55. 5-12 Jul 1830 [’Chamaelaucieae’]; Myrtineae Burnett, Outl. Bot.: 616, 617, 1092, 1128. Jun 1835; Myrrhiniaceae Arn. in Ann. Nat. Hist. 3: 154. 1839 [’Olinieae, or Myrrhinieae’]; Heteropyxidaceae Engl. et Gilg, Syllabus, ed. 8: 281. Jan-Feb 1920, nom. cons.; Kaniaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 245. 20 Jul 1943; Psiloxylaceae Croizat, Principia Bot.: 604, 1161, 1164. Jun 1961

Genera/species 130–131/5.390–5.460

Distribution Tropical, subtropical and warm-temperate regions on both hemispheres, with their largest diversity in Australia and South America.

Fossils Myrtaceae are often found as fossils in the Cenozoic of North America and Europe and fossil leaves and pollen grains are frequently recorded from the Australian Cenozoic. Myrtaceidites comprises pollen grains, possibly of Myrtaceae, from the Maastrichtian of Australia and the Paleocene of New Zealand. Paleomyrtinaea, a probably quinquelocular fruit with parietal placentation and C-shaped embryos, has been described from the Late Paleocene of North Dakota and the Early Eocene of British Columbia. Fossils assigned to Eucalyptus have been found in Early Eocene strata in Patagonia (Argentina).

Habit Usually bisexual (rarely andromonoecious, polygamomonoecious, dioecious, or androdioecious), usually evergreen (rarely deciduous) trees or shrubs. Often aromatic. Bark often exfoliating. Numerous species are xerophytes. Osbornia consists of mangrove trees.

Vegetative anatomy Ectomycorrhiza usually present (sometimes also endomycorrhiza; in Eucalyptus arbuscular mycorrhiza). Phellogen ab initio superficial or pericyclic. Cork tissue usually stratified. Polyderm present. Primary vascular tissue usually bicollateral. Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements tracheids or fibre tracheids, with simple or bordered pits, usually non-septate (rarely septate; sometimes also vasicentric). Wood rays uniseriate or multiseriate, heterocellular or homocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, often banded, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, unilateral, or banded. Tyloses sometimes frequent. Intraxylary phloem usually present. Secondary phloem in young stems usually stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids Ss type; sieve tubes usually with non-dispersive protein bodies. Nodes 1:?, unilacunar with ? leaf traces. Sclerenchymatous idioblasts abundant. Schizolysigenous secretory cavities (in, e.g., medulla and/or cortex) with ethereal oils. Heartwood often with gum-like substances. Wood rays with or without silica bodies. Calciumoxalate as prismatic or acicular crystals, styloids, crystal sand or other types of crystals often abundant.

Trichomes Hairs usually unicellular, uniseriate, simple (sometimes bi- or multicellular, furcate); bristle-like glands present or absent.

Leaves Usually opposite (rarely alternate, spiral, or verticillate), simple, entire, often coriaceous, with ? ptyxis. Stipules rudimentary, modified into colleters or hairs, or absent; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate or parallelodromous (rarely palmate; leaves sometimes one-veined). Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids as platelets, chemically characterized by presence of primary alcohols and triterpenoids, or as tubuli dominated by β-diketones, or as parallel grouped (usually non-entire) platelets (Hypericum type). Domatia in Heteropyxis as pockets. Lamina with gland-dots and schizogenous secretory cavities with ethereal oils. Sclerenchymatous idioblasts abundant. Epidermis with or without mucilaginous idioblasts. Leaf margin entire. Extrafloral nectaries rarely present in leaf axils.

Inflorescence Terminal or axillary, usually paniculate (rarely racemose, spicate, capitate or corymbose; flowers rarely solitary, axillary). Inflorescence sometimes pseudanthium with involucre of showy bracts.

Flowers Usually actinomorphic (rarely zygomorphic, e.g. in Calothamnus). Hypanthium present. Usually epigyny or half epigyny (ovary adnate to receptacle entirely or only in lower part; rarely almost hypogyny; in Heteropyxis and Psiloxylon hypogyny). Sepals (three or) four or five (to eight), with usually imbricate quinquncial (sometimes valvate) aestivation, free (sometimes reduced or modified into caducous circumscissile calyptra; sepals in Verticordia fringed). Petals (three or) four or five (to twelve), with imbricate aestivation, often early caducous, clawed, free or connate at base (sometimes modified into caducous circumscissile calyptra or absent). Nectariferous disc inserted on apex of ovary or on prolonged hypanthium. Secondary pollen display sometimes present.

Androecium Stamens usually c. 20 to more than 150 (sometimes eight to ten, rarely four or five), in one or several whorls, alternisepalous or antesepalous, alternipetalous or antepetalous, basically antepetalous. Filaments usually inflexed in bud (sometimes upright or twice folded), free or often connate at base into four or five usually antepetalous fascicles, or connate into tube, free from tepals, inserted on hypanthium, often very long; centripetally or centrifugally developing. Anthers dorsifixed, versatile, usually tetrasporangiate (rarely monosporangiate or trisporangiate), introrse, latrorse or extrorse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by pores); connective usually with one or several terpene-producing apical oil-glands. Tapetum secretory. Staminodia usually absent (present in female flowers of Psiloxylon).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually markedly triangular in polar view, (2–)3(–4)-colporate or (2–)3(–4)-porate (rarely (2–)3(–4)-colpate), often (seemingly) syncolporate, not heterocolpate (pseudocolpi absent), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate or scabrate to psilate.

Gynoecium Pistil composed of usually two to five (rarely up to 18) connate antesepalous or antepetalous carpels, or odd carpel abaxial. Ovary usually inferior or semi-inferior, bilocular to quinquelocular (rarely up to 18-locular or unilocular and pseudomonomerous). Style single, sunken, usually elongate, persistent, simple or branched in upper part. Stigma usually single, capitate, punctate, trilobate or quadrilobate (rarely peltate; stigmas sometimes three or four), Dry or Wet type. Male flowers often with pistillodium.

Ovules Placentation usually axile (rarely parietal to basal, when ovary unilocular). Ovules usually c. 30 to more than 150 (rarely one to 15) per carpel, anatropous, hemianatropous or campylotropous, ascending, usually bitegmic (rarely unitegmic), crassinucellar. Micropyle bistomal or endostomal (sometimes exostomal?). Outer integument two to six (to twelve) cell layers thick. Inner integument two to four cell layers thick. Obturator sometimes present. Hypostase absent. Parietal tissue two to twelve cell layers thick. Nucellar cap absent. Megagametophyte usually monosporous, Polygonum type (in Heteropyxis and Psiloxylon disporous, Allium type). Synergids often with a filiform apparatus. Antipodal cells sometimes not developing (with nuclei early degenerating). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad (or adventitious). Polyembryony frequent in several genera.

Fruit Usually a loculicidal capsule or a berry (rarely a septicidal or denticidal capsule or a pyxidium, drupe, schizocarp, or nutlike fruit). Hypanthium usually accrescent in fruit.

Seeds Aril absent. Operculum often? present. Seed coat testal. Testa sometimes winged, often with crystalliferous layers, sometimes with endotestal micropylar plug, sometimes multiplicative, often partially sclerotic. Exotesta and/or endotesta often thickened. Exotegmen usually absent or collapsed. Endotegmen often collapsed. Perisperm not developed. Endosperm usually absent or scarce (with oils and starch). Embryo straight, curved or spirally twisted, well differentiated, with or without chlorophyll. Cotyledons two, accumbent or incumbent, often plicate, often connate. Germination phanerocotylar or cryptocotylar.

Cytology n = (5–)11, (12, 22)

DNA Plastid gene infA lost/defunct (Callistemon).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, delphinidin, ethereal oils (triterpenes etc.), ursolic acid, ellagic and gallic acids, piperidine, pyrrolidine and pyrrolizidine alkaloids (polyhydroxyalkaloids, usually frequent), saponins, and phenylalanine-derived cyanogenic compounds present. Many species produce gum. Raffinose and stachyose in phloem exudates. Aluminium accumulated in some species.

Use Ornamental plants, fruits (Psidium, Feijoa, Campomanesia, Eugenia, etc.), spices (Eucalyptus, Pimenta, Syzygium), gums, essential oils for perfumes, medicinal plants (Eucalyptus), timber, paper pulp (Eucalyptus), drainage of swamps (Eucalyptus, Metrosideros).

Systematics Myrtaceae are sister to Vochysiaceae.

The clade [Heteropyxis+Psiloxylon] is sister to the remaining Myrtaceae.

Psiloxyloideae (Croizat) Schmid in Taxon 29: 559. 14 Nov 1980

2/4. Central and southeastern Africa, the Mascarene Islands. Dioecious trees or shrubs. Leaves alternate (spiral). Petals clawed. Stamens erect in bud, not fasciculated, with separate leaf traces. Anthers with thecae separately dehiscing. Female flowers with staminodia. Pistil composed of (two or) three connate carpels. Ovary superior, with narrow base. Male flowers with pistillodium. Megagametophyte disporous, 8-nucleate, Allium type. Endotestal cells periclinally elongate, crystalliferous. x = 12. Tannins abundant or absent.

Heteropyxis

1/3. Heteropyxis (3; central and southeastern Africa). – Usually dioecious (sometimes monoecious), deciduous aromatic trees or shrubs. Cork tissue not stratified. Pits bordered. Vasicentric tracheids absent. Axial (apotracheal) xylem parenchyma absemt. Fibres non-septate. Crystalloids as platelets; rhomboidal crystals only in wood ray cells. Leaves simple, entire. Stipules minute. Cuticular wax crystalloids as small platelets. Lamina with domatia and superepidermal secretory cavities containing ethereal oils. Inflorescence terminal panicle. Flowers actinomorphic, small. Hypanthium present. Hypogyny. Sepals (four or) five, persistent. Petals (four or) five, gland-dotted, caducous, free, alternisepalous, inserted on inner margin of hypanthium. Nectariferous disc present in lower part of hypanthium. Stamens (four to) five to eight (to ten), antesepalous (sometimes also one, two or three antepetalous), usually obdiplostemonous, erect in bud. Filaments free from each other and from tepals, not in fascicles. Anthers dorsifixed, versatile?, tetrasporangiate, usually latrorse (rarely introrse), longicidal (dehiscing by longitudinal slits); connective with one to three secretory cavities. Female flowers with (four or) five to eight (to ten) staminodia. Pollen grains resembling those in Myrtoideae, syncolporate, with depressions between colpi, not heterocolpate (pseudocolpi absent). Pistil composed of (two or) three connate carpels. Ovary bilocular (or trilocular). Style sunken into ovary. Stigma capitate, persistent. Placenta axile. Ovules numerous, hemianatropous. Fruit a loculicidal capsule (partially enclosed by persistent calyx). Testa with apical wing (sometimes one at micropyle). Exotesta with tangentially elongate cells with scalariform to reticulately thickened walls. Exotegmic cells elongate. Endosperm absent. Embryo straight. Terpenes not found.

Psiloxylon

1/1. Psiloxylon (1; P. mauritianum; Mauritius, Réunion). – Monoecious or dioecious (rarely polygamodioecious), evergreen tree with white bark. Phellogen ab initio pericyclic. Young stem with secretory canals. Intraxylary phloem present. Secondary lateral growth from normal cylindrical cambium. Xylem without fibre tracheids. Axial parenchyma usually scarce paratracheal. Wood fibres septated, with rhomboidal crystals. Hairs absent. Leaves seemingly alternate, simple, entire. Stipules modified into colleters; leaf sheath absent. Lamina gland-dotted, with schizogenous secretory cavities not containing ethereal oils. Inflorescence axillary panicle. Flowers actinomorphic. Pedicel articulated. Hypanthium present. Hypogyny. Sepals four or five (or six), with imbricate aestivation, coriaceous, persistent, free. Petals four or five (or six), with imbricate aestivation, shortly clawed, coriaceous, caducous, punctate, free. Stamens (4+4 or) 5+5 (or 6+6), twice as many as petals, diplostemonous, erect in bud. Filaments inserted at perigynous nectariferous disc, free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Female flowers with staminodia. Pollen grains tricolporate, parasyncolporate, ?-cellular at dispersal. Exine rugulate. Pistil composed of three or four connate carpels. Ovary trilocular (or quadrilocular), often stipitate. Style very short, sunken into ovary, or absent. Stigma large, fleshy, usually trilobate (sometimes bilobate or quadrilobate), persistent in fruit. Male flowers with pistillodia. Placentation axile. Ovules c. 30 to c. 70 per carpel, anatropous or hemicampylotropous. Fruit a many-seeded berry, gland-dotted. Exotestal cells large. Exotegmen crushed. Endosperm absent. Embryo well differentiated, fleshy, straight. Cotyledons two. Ethereal oils and polyhydroxyalkaloids present.

Myrtoideae Sweet, Fl. Australas.: ad t. 10. 1 Aug 1827 [‘Myrteae’]

128–129/5.390–5.460. Trees or shrubs. Ectomycorrhiza often present (in species with dry fruits). Sieve tubes with non-dispersive protein bodies. Hairs usually unicellular (sometimes multicellular). Leaves usually opposite (sometimes alternate), with various ptyxis. Stipules two or several, often modified into colleters, or absent. Venation usually pinnate (rarely palmate). Stomata sometimes paracytic. Silica sometimes present in leaf tissues. Hypanthium present (often operculate and caducous). Usually epigyny (rarely hypogyny to half epigyny). Perianth undifferentiated in some genera closely allied to Eucalyptus. Calyx or entire perianth sometimes calyptrate, as caducous lid. Petals (absent to) four or five (to twelve), often caducous. Stamens numerous, large, very varying in number and shape (rarely five, as many as petals or in antepetalous or antesepalous fascicles; rarely ten), principally antepetalous. Pollen grains often syncolpate. Pistil composed of two (to 18) connate alternipetalous or antepetalous carpels (sometimes with odd carpel abaxial). Ovary usually inferior, with wide base, sometimes unilocular. Placentae thick. Transseptal vascular bundles present. Style without small vascular bundles. Stigma punctate to capitate (rarely peltate), Dry or Wet type. Placentation rarely basal. Ovules one to four (to ten) per carpel, usually bitegmic (rarely unitegmic). Usually a capsule (rarely a berry). Testa multiplicative, more or less sclerotic or exotesta thickened (exotesta in species with capsule, sclerotised testa in species with berry). Endotesta often thickened. Sclerotic palisade cells sometimes present at micropyle. Embryo in Eugenia undifferentiated. Polyhydroxy-alkaloids often abundant. Many species gum-producing. – The phylogeny of Myrtoideae is largely unresolved (Wilson & al. 2005). The clade [Xanthostemoneae+Lophostemoneae] is sister group to the remaining Myrtoideae. Syzygieae and Tristaneae form a monophletic group. Lindsayomyrteae are probably sister to [Chamelaucieae+Leptospermeae]. Myrteae and Syzygieae have predominant baccate fruit and seeds with sclerotic testa.

[Xanthostemoneae+Lophostemoneae]

Xanthostemoneae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 14. 16 Mar 2005

3/c 50. Pleurocalyptus (3; New Caledonia), Purpureostemon (1; P. ciliatus; New Caledonia), Xanthostemon (c 45; the Philippines, East Malesia to New Guinea, northern Australia to eastern Queensland, the Solomon Islands, New Caledonia). – Central and East Malesia to New Guinea, northern Australia to eastern Queensland, the Solomon Islands, New Caledonia.

Lophostemoneae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 14. 16 Mar 2005

4/7–9. Kjellbergiodendron (1–3; K. celebicum; Sulawesi, New Guinea), Lophostemon (4; southern New Guinea, northern and eastern Australia), Welchiodendron (1; W. longivalve; New Guinea, northern Queensland), Whiteodendron (1; W. moultonianum; Borneo). – Borneo, Sulawesi, New Guinea, northern and eastern Australia.

The remaining Myrtoideae

Osbornieae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 14. 16 Mar 2005

1/1. Osbornia (1; O. octodonta; coasts in the Philippines, East Malesia to New Guinea, northern Australia to eastern Queensland, Palau). – Evergreen mangrove tree or shrub without pneumatophores.

Melaleuceae Burnett, Outlines Bot.: 712. Feb 1835

10/c 320. Beaufortia (19; southwestern Western Australia), Callistemon (c 30; Australia, Tasmania, New Caledonia), Calothamnus (c 25; southwestern Western Australia), Conothamnus (3; southwestern Western Australia), Eremaea (16; southwestern Western Australia),Lamarchea (2; Western Australia), Melaleuca (c 250; Australia, Tasmania, New Caledonia, few species in tropical Asia and eastwards to islands in the Pacific), Petraeomyrtus (1; P. punicea; northern Northern Territory), Phymatocarpus (3; southwestern Western Australia), Regelia (5–6; southwestern Western Australia). – Tropical Asia and eastwards to islands in the Pacific, Australia, Tasmania, New Caledonia, with their highest diversity in Australia.

Kania clade

8/c 55. Barongia (1; B. lophandra; northeastern Queensland), Basisperma (1; B. lanceolata; New Guinea), Kania (6; the Philippines, New Guinea), Lysicarpus (1; L. angustifolius; eastern Queensland), Mitrantia (1; M. bilocularis; northeastern Queensland), Ristantia (3; northeastern Queensland), Sphaerantia (2; northeastern Queensland), Tristaniopsis (c 40; Southeast Asia, Malesia to New Guinea, eastern Australia). – Southeast Asia, Malesia to New Guinea, eastern Australia.

Backhousieae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 15. 16 Mar 2005

1–2/10. Backhousia (8; eastern Queensland, eastern New South Wales), Choricarpia (2; eastern Queensland, eastern New South Wales; in Backhousia?). – Eastern Queensland, eastern New South Wales.

Metrosidereae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 15. 16 Mar 2005.

2/55–60. 'Metrosideros’ (55–60; East Malesia to New Guinea, northeastern Queensland, eastern New South Wales, the Solomon Islands, New Caledonia, Lord Howe, New Zealand, Fiji, the Bonin Islands, the Tuamotus, the Hawaiian Islands, one species, M. angustifolia, in Western Cape in South Africa; polyphyletic), Tepualia (1; T. stipularis; southern Chile, southern Argentina). – South Africa, East Malesia, eastern Australia, Melanesia, New Zealand, Polynesia, temperate South America.

[Syzygieae+Tristanieae]

Syzygieae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 15. 16 Mar 2005

2/c 1.200. Anetholea (1; A. anisata; New South Wales), Syzygium (c 1.200; southeastern Africa, Madagascar, tropical and subtropical Asia, northern and eastern Australia, Lord Howe, and eastwards to islands in the Pacific to Samoa). – Tropical and subtropical regions in the Old World to the Pacific.

Tristanieae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 15. 16 Mar 2005

3/28. Thaleropia (3; New Guinea, northeastern Queensland), Tristania (1; T. neriifolia; southeastern Queensland, eastern New South Wales),Xanthomyrtus (24; Borneo, the Philippines, Sulawesi, the Moluccas, New Guinea, the Bismarck Archipelago, New Caledonia). – Malesia to New Guinea, eastern Queensland, eastern New South Wales, New Caledonia.

Myrteae (A. P. de Candolle) DC. in D. F. L. von Schlechtendal in Linnaea 2: 504. Jul 1827

c 50/2.165–2.225. Acca (3; South America), Accara (1; A. elegans; Brazil), Algrizea (1; A. macrochlamys; Bahia in Brazil), Amomyrtella (1; A. quili; northern Argentina), Amomyrtus (2; Chile), Archirhodomyrtus (6; eastern Queensland, northeastern New South Wales, New Caledonia), Austromyrtus (3; southeastern Queensland, eastern New South Wales), Blepharocalyx (3; tropical and subtropical South America), Calycolpus (14; tropical America), Calycorectes (17; tropical America), Calyptranthes (c 130; tropical America; in Myrcia?), Calyptrogenia (6; tropical America), Campomanesia (c 80; South America), Chamguava (3; Central America), Curitiba (1; C. prismatica; Planalto in southeastern Brazil), Decaspermum (c 30; tropical Asia and eastwards to New Guinea, eastern Queensland, northeastern New South Wales and islands in western Pacific), Eugenia (950–1.000; tropical regions on both hemispheres, with their highest diversity in tropical America), Gossia (20; New Guinea, eastern Queensland, eastern New South Wales, New Caledonia), Kanakomyrtus (6; New Caledonia), Legrandia (1; L. concinna; Chile), Lenwebbia (2; northeastern Queensland, eastern New South Wales), Lithomyrtus (11; northern Australia), Lophomyrtus (2; New Zealand; incl. Neomyrtus?), Luma (2; Peru, Chile, Argentina), Marlierea (c 60; tropical South America; in Myrcia?), Meteoromyrtus (1; M. wynaadensis; southern India), Mitranthes (11; tropical America), Mosiera (3–4; Central America), Myrceugenia (c 40; southeastern Brazil, Chile, Juan Fernandez, Argentina), Myrcia (c 450; tropical America, with their largest diversity in southeastern Brazil; incl. Calyptranthes?, Marlierea?), Myrcianthes (c 30; tropical America), Myrciaria (c 60; tropical America; incl. Paramyrciaria?), Myrrhinium (1; M. atropurpureum; tropical South America), Myrtastrum (1; M. rufopunctatum; New Caledonia), Myrtella (2; New Guinea, one species, M. bennigseniana, extending to Guam), Myrteola (3; southern South America, tropical American mountains), Myrtus (2; the Mediterranean, North Africa), Neomyrtus (1; N. pedunculata; New Zealand, Stewart Island; in Lophomyrtus?), Octamyrtus (6; the Moluccas, New Guinea), Paramyrciaria (6; South America; in Myrciaria?), Pilidiostigma (6; eastern Queensland, northeastern New South Wales), Pimenta (15; tropical America), Plinia (25–30; tropical America), Psidium (c 70; tropical America), Rhodamnia (28; Southeast Asia, Hainan, Malesia, New Guinea, northern and eastern Australia, the Solomon Islands, New Caledonia), Rhodomyrtus (11; tropical Asia, Malesia to New Guinea, eastern Queensland, eastern New South Wales, New Caledonia), Siphoneugena (8; tropical America), Stereocaryum (3; New Caledonia), Ugni (4; tropical and subtropical America), Uromyrtus (15–20; Borneo to New Guinea, eastern Queensland, northeastern New South Wales, New Caledonia). – Pantropical, with their highest diversity in tropical America.

Eucalypteae (Benth. et Hook. f.) Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 16. 16 Mar 2005

7/>835. Allosyncarpia (1; A. ternata; Arnhem Land in Northern Territory),Angophora (15; eastern Australia), Arillastrum (1; A. gummiferum; southern New Caledonia), Corymbia (c 115; southern New Guinea, Australia, Tasmania), Eucalyptopsis (2; Buru, New Guinea), Eucalyptus (>700; Mindanao, Sulawesi, East Malesia to New Guinea, Australia, Tasmania), Stockwellia (1; S. quadrifida; Atherton Tableland in northeastern Queensland). – The Philippines, Sulawesi, the Moluccas, New Guinea, Australia, Tasmania, New Caledonia.

Syncarpieae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 16. 16 Mar 2005

1/3. Syncarpia (3; eastern Queensland, eastern New South Wales). – Trees.

[Lindsayomyrteae+[Leptospermeae+Chamelaucieae]]

Lindsayomyrteae Peter G. Wilson in P. G. Wilson et al., Plant Syst. Evol. 251: 16. 16 Mar 2005

1/1. Lindsayomyrtus (1; L. racemoides; the Moluccas, northeastern Queensland). – Tree.

[Leptospermeae+Chamelaucieae]

Leptospermeae (A. P. de Candolle) DC. in D. F. L. von Schlechtendal in Linnaea 2: 504. Jul 1827

c 8/145–150. Agonis (12; southwestern Western Australia), Angasomyrtus (1; A. salina; southwestern Western Australia), Asteromyrtus (7; southern New Guinea, northern Australia), Homalospermum (1; H. firmum; coasts in southwestern Western Australia), Kunzea (c 35; southwestern Western Australia, southeastern Australia, Tasmania, one species, K. ericoides, in New Zealand), ’Leptospermum’ (c 85; Southeast Asia, Malesia to New Guinea, Australia, Tasmania, New Zealand; polyphyletic),Neofabricia (3; northern Queensland), Pericalymma (3; southwestern Western Australia). – Southeast Asia, Malesia to New Guinea, Australia, Tasmania, New Zealand.

Chamelaucieae (A. P. de Candolle) DC. in D. F. L. von Schlechtendal in Linnaea 2: 504. Jul 1827 [‘Chamaelaucieae’]

24/495–500. Actinodium (3; southwestern Western Australia), Aluta (5; western and central Australia), Astartea (8; southwestern Western Australia), Babingtonia (16; Malesia to New Guinea, southwestern Western Australia, southeastern South Australia, western Victoria, New Caledonia), Baeckea (14; Australia, Tasmania, New Caledonia, one species, B. frutescens, extending to Southeast Asia, China), Balaustion (2; southwestern Western Australia), Calytrix (c 75; Australia, Tasmania, with their highest diversity in southwestern Western Australia), Chamelaucium (23; southwestern Western Australia), Corynanthera (1; C. flava; southwestern Western Australia), Darwinia (c 45; southern Western Australia, South Australia, Victoria, southern New South Wales), Harmogia (14; southwestern Western Australia, southeastern Queensland, eastern New South Wales), Homalocalyx (11; Western Australia, Arnhem Land in Northern Territory, southern Queensland), Homoranthus (28; eastern Queensland, eastern New South Wales, southern South Australia), Hypocalymma (c 30; southwestern Western Australia), Kardomia (6; southeastern Queensland, northeastern New South Wales), Malleostemon (6; southwestern Western Australia), Micromyrtus (22; Australia), Ochrosperma (6; southeastern Queensland, eastern New South Wales), Pileanthus (8; southwestern Western Australia), Rinzia (12; southwestern Western Australia), Sannantha (16; eastern Queensland, eastern New South Wales, eastern Victoria), Scholtzia (c 15; western and southwestern Western Australia), Stenostegia (1; S. congesta; northern Northern Territory; in Baeckea?), Thryptomene (c 30; southern, central and northeastern Australia, Tasmania), Verticordia (c 100; Western Australia, northern Northern Territory, with their highest diversity in southwestern Western Australia). – Southeast Asia, southern China, Malesia to New Guinea, Australia, Tasmania, New Caledonia, with their largest diversity in southwestern Western Australia.

Unplaced Myrtoideae

Cloezia (8; New Caledonia), Hottea (5; Hispaniola), Seorsus (4; Borneo, Northern Territory, southwestern Western Australia).

Cladogram (simplified) of Myrtaceae based on DNA sequence data (Wilson & al. 2005). There is weak support for Osbornieae being sister-group to Melaleuceae.

OLINIACEAE Arn.

( Back to Myrtales )

Arnott in Ann. Nat. Hist. 3: 154. 1839 [’Olinieae or Myrrhinieae’], nom. cons.

Plectroniaceae Hiern, Cat. Afr. Pl. Welw. 1(2): 369. Mar 1898

Genera/species 1/8–10

Distribution Southern and eastern Africa.

Fossils Unknown.

Habit Bisexual, evergreen shrubs. Young stems and branches quadrangular in cross-section or terete. Inner bark usually almond-scented.

Vegetative anatomy Phellogen ab initio subepidermal. Primary medullary rays narrow. Vessel elements diffuse, with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple pits, septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal scanty (apotracheal parenchyma absent). Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace. Intraxylary phloem present. Cortex with branched sclereids. Calciumoxalate raphides absent.

Trichomes Hairs unicellular or absent; glandular hairs present or absent.

Leaves Opposite or verticillate, simple, entire, coriaceous, with ? ptyxis. Stipules very small, rudimentary, cauline or inserted at leaf base, or absent; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate; midvein with apical swelling or glandular tip. Stomata paracytic or anomocytic (or cyclocytic). Cuticular wax crystalloids? Domatia as pits or absent. Terminal sclereids present. Mesophyll with calciumoxalate as solitary prismatic crystals. Leaf margin entire.

Inflorescence Terminal or axillary, thyrsoid-paniculate.

Flowers Actinomorphic, small. Hypanthium tubular, with (four or) five apical teeth (representing epicalyx? or calyx?). Epigyny. Sepals (four or) five, with open (imbricate?) aestivation, petaloid, spatulate, alternate with hypanthial teeth, caducous, free. Petals (four or) five, with valvate aestivation, scale-like, concave, thick, hairy, inserted on adaxial side of hypanthial margin, free. Nectariferous disc absent. Idioblasts containing calciumoxalate druses abundant.

Androecium Stamens (four or) five, obhaplostemonous, alternisepalous, antepetalous. Filaments short, inserted below petals on adaxial side of hypanthial margin, inflexed in bud, free from each other and from tepals. Anthers dorsifixed (basifixed?), non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits, microsporogenous tissue often divided into two packages by band of tapetal cells); connective distally abaxially expanded; endothecium ephemeral. Tapetum secretory, with binucleate cells. Staminodia four to ten, in one or two whorls (one whorl sometimes rudimentary), extrastaminal, often scale-like or petaloid.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, heteropolar (with one pole colpate and one pole heterocolpate, with pseudocolpi limited to one of two poles, half pseudocolpi), shed as monads, bicellular at dispersal; colpi asymmetrical: each colpus consisting of one long segment on larger polar surface and one short segment on smaller polar surface. Exine tectate, with columellate infratectum, psilate.

Gynoecium Pistil composed of (two to) five connate antesepalous carpels. Ovary inferior, usually quinquelobate (rarely bilobate to quadrilobate). Transseptal vascular bundles present. Style single, simple, subulate. Stigma capitate to clavate, papillate (sometimes commissural), Wet type? Pistillodium absent?

Ovules Placentation axile. Ovules two to ten per carpel, campylotropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument approx. five cell layers thick. Inner integument ? cell layers thick. Hypostase present. Parietal tissue three or four cell layers thick. Chalaza strongly vascularized. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A single- or several-seeded drupe.

Seeds Aril absent. Testa thick, three- or four-layered, vascularized. Exotegmen fibrous. Endotegmen? Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, without suspensor, chlorophyll? Cotyledons two, spirally twisted or irregularly folded. Germination?

Cytology n = 12, c. 15, c. 20 – Polyploidy probably occurring.

DNA

Phytochemistry Insufficiently known. Non-hydrolyzable tannins and phenylalanine-derived cyanogenic compounds (prunasine, an almond-smelling cyanogenic glycoside) present. Aluminium not accumulated.

Use Medicinal plants, timber.

Systematics Olinia (8–10; South Africa to Ethiopia and Angola).

Olinia is sister to Penaeaceae.

ONAGRACEAE Juss.

( Back to Myrtales )

de Jussieu, Gen. Plant.: 317. 4 Aug 1789 [’Onagrae’], nom. cons.

Epilobiaceae Vent., Tabl. Règne Vég. 3: 307. 5 Mai 1799 [‘Epilobianae’]; Oenotheraceae C. C. Robin, Voy. Int. Louisiane 3: 489. 14-21 Dec 1807 [’Oenothera’]; Circaeaceae Bercht. et J. Presl, Přir. Rostlin: 232. Jan-Apr 1820 [‘Circeaceae’]; Isnardiaceae Martinov, Tekhno-Bot. Slovar: 348. 3 Aug 1820 [’Isnardiae’]; Jussiaeaceae Martinov, Tekhno-Bot. Slovar: 350. 3 Aug 1820 [’Jussieae’]; Onagrales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 232. Jan-Apr 1820 [‘Onagrae’]; Circaeales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 48. Sep-Oct 1835 [‘Circaeaceae’]; Epilobiales Vent. in C. F. P. von Martius, Consp. Regn. Veg.: 48. Sep-Oct 1835 [’Epilobiaceae’]; Oenotherales Bromhead in Edinburgh New Philos. J. 24: 409. Apr 1838; Onagrineae Rchb., Deutsch. Bot. Herb.-Buch: lxviii. Jul 1841; Oenotheropsida Brongn., Enum. Plant. Mus. Paris: xxx, 117. 12 Aug 1843 [’Oenotherineae’]; Fuchsiaceae (DC.) Lilja, Skånes Fl., ed. 2: 846, 980. Apr-Dec 1870; Lopeziaceae (Spach) Lilja, Skånes Fl., ed. 2: 980. Apr-Dec 1870

Genera/species 22/655–660

Distribution: Cosmopolitan except Antarctica, with their largest diversity in southwestern North America.

Fossils Seeds of Ludwigia have been found in Oligocene layers onwards in Europe and Asia, and Fuchsia pollen is recorded from the Eocene of Argentina. Seeds of Palaeeucharidium from the Eocene London Clay, southern England, may be a representative of Onagraceae.

Habit Usually bisexual (rarely unisexual), usually perennial, biennial or annual herbs (rarely trees or shrubs, e.g. in Fuchsia). Some species are aquatic.

Vegetative anatomy Phellogen ab initio deeply seated. Polyderm present. Endodermis sometimes prominent. Primary vascular tissue usually bicollateral. Secondary lateral growth normal or anomalous (from concentric cambia or from cylindrical cambium). Vessel elements with simple perforation plates; lateral pits alternate, usually bordered (sometimes simple) pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal banded, or paratracheal scanty vasicentric, or absent. Tyloses sometimes frequent. Interxylary phloem often present. Sieve tube plastids Ss type. Nodes 1:1, unilacunar with one leaf trace. Epidermis often with cells containing ethereal oils. Calciumoxalate raphides, prismatic crystals and styloids often present.

Trichomes Hairs unicellular or multicellular simple (sometimes with widened base); glandular hairs often frequent.

Leaves Usually opposite (sometimes alternate, rarely verticillate), simple, usually entire (rarely lobed), with involute to flat ptyxis. Stipules usually small, intrapetiolar, caducous, or absent (in Ludwigia often conspicuous); leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata with three or more subsidiary cells, anomocytic, anisocytic, tetracytic or cyclocytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type) or transversely ridged rodlets, chemically dominated by ketones. Epidermis with or without mucilaginous idioblasts. Mesophyll without sclerenchymatous idioblasts. Calciumoxalate raphides abundant. Leaf margin serrate or entire.

Inflorescence Usually terminal (rarely axillary), panicle (rarely cyme), raceme or spike, or flowers solitary axillary. Floral prophylls (bracteoles) sometimes absent.

Flowers Usually actinomorphic (sometimes zygomorphic). Hypanthium usually present (absent in Ludwigia). Epigyny or half epigyny. Sepals (two to) four (to seven), with valvate aestivation, inserted on margin of hypanthium, free, usually caducous. Petals (two to) four (to seven; in Circaea two; in Lopezia two adaxial petals recurved, with pseudonectaries on claws), with imbricate, valvate or contorted aestivation, inserted on margin of hypanthium, often clawed (rarely absent), free, caducous. Nectaries present on adaxial side of hypanthium. Disc intrastaminal, annular, on top of ovary, or absent.

Androecium Stamens usually 4+4 or 6+6 (in Lopezia one fertile and one sterile stamen; in Circaea two antesepalous stamens). Filaments free from each other and from tepals, inserted on adaxial side of hypanthium or around epigynous nectariferous disc. Anthers dorsifixed, versatile, tetrasporangiate or polythecate, usually introrse (in Lopezia extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia one to four (in Lopezia one petaloid), or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually strongly triangular, (2–)3(–6)-pororate, (2–)3(–6)-colpate or (2–)3(–6)-colporate, with bulging apertures, starchy, usually covered by viscin threads of various types (annular-vermiform, moniliform, smooth, etc.), not heterocolpate (pseudocolpi absent), shed as monads, tetrads or polyads, bicellular at dispersal. Exine tectate, with columellate or acolumellate infratectum, verrucate, granulate or smooth. Ectexine with paracrystalline bodies.

Gynoecium Pistil composed of (two to) four (to seven) connate alternisepalous carpels. Ovary inferior or semi-inferior, (unilocular to) quadrilocular (to septalocular). Style single, simple. Stigma single, capitate to quadrilobate (rarely commissural), papillate or non-papillate, Dry or Wet type. Male flowers often with pistillodium.

Ovules Placentation usually axile (rarely parietal). Ovules usually few to many (rarely one or two) per carpel, anatropous, pendulous or ascending, bitegmic, crassinucellar. Micropyle bistomal. Outer integument two to five cell layers thick. Inner integument two or three cell layers thick. Hypostase usually present. Parietal tissue approx. two (to c. 25) cell layers thick. Nucellar cap approx. two cell layers thick. Archespore often multicellular. Megagametophyte quadrinucleate, Oenothera type, with micropylar megaspore functional. Synergids with a filiform apparatus. Antipodal cells not developed. Endosperm development ab initio nuclear; endosperm diploid. Endosperm haustoria? Embryogenesis onagrad.

Fruit Usually a loculicidal (sometimes also septicidal) capsule dehiscing from apex downwards (in Fuchsia a berry; in Circaea and species of Oenothera a nutlet).

Seeds Aril absent. Seed coat endotestal. Exotesta often densely hairy to papillate, with inner cell walls thickened and lignified. Mesotestal cells sometimes thickened, sclerotic. Endotestal cells stellate. Exotegmen often fibrous. Endotegmic cells sometimes longitudinally elongate, tanniniferous, with inner walls thickened. Perisperm not developed. Endosperm absent. Embryo straight or almost straight, well differentiated, oily, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology x = 7, 9–11, 18 – Permanent reciprocal chromosomal translocations occurring in many Onagreae (e.g. species of Oenothera, chromosomes forming ring due to permanent translocations, entire genome inherited as one unit).

DNA Inversion of 45 kb present in plastid genome in Oenothera sect. Oenothera. Plastid gene clpP lost at least in Oenothera missouriensis. Plastid gene infA lost in Oenothera.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavonoid sulphates, delphinidin, ursolic acid, ellagic acid, alkaloids, and cyanogenic compounds present. Saponins not found. Raffinose and stachyose present in phloem exudates. Dissolved oxalates accumulated.

Use Ornamental plants, seed oils for medicinal purpose.

Systematics Onagraceae are sister-group to Lythraceae.

Ludwigia is sister to the remaining Onagraceae.

Jussiaeoideae Beilschm. in Flora 16(Beibl. 7): 96, 108. 14 Jun 1833 [‘Jussiaceae’]

1/80–85. Ludwigia (80–85; cosmopolitan, with their largest diversity in America). – Stipules often prominent. Hypanthium absent. Flowers tetramerous or pentamerous. Pollen grains usually shed as tetrads (rarely monads). Nectary present on apex of ovary. Style short, with small vascular bundles. Stigma capitate. Hypostase present. Parietal tissue three to six cell layers thick. Megasporocyte single.

Onagroideae Beilschm. in Flora 16(Beibl. 7): 96. 14 Jun 1833 [‘Onagrariae’]

21/c 575. Hauya (2; Central America); Circaea (7; temperate regions on the Northern Hemisphere), Fuchsia (c 105; New Zealand, Tahiti, Central and South America); Lopezia (22; Mexico, Central America), Megacorax (1; M. gracielanus; Durango in Mexico); Gongylocarpus (2; Mexico, Central America); Epilobium (c 165; temperate and arctic-alpine regions on both hemispheres, tropical mountains, including Tasmania, New Zealand and adjacent islands, Tierra del Fuego, with their highest diversity in western North America), Chamaenerion (8; temperate regions on the Northern Hemisphere); Xylonagra (1; X. arborea; Baja California in northwestern Mexico); Taraxia (c 10; North America, Mexico), Clarkia (>40; western North America, Mexico, southern South America), Gayophytum (9; western North America, western South America), Chylismiella (1; C. pterosperma; southwestern United States); Eulobus (4; California, Arizona, Baja California in northwestern Mexico), Oenothera (c 145; northwestern Europe, America, with their highest diversity in western North America), Chylismia (16; western North America, northwestern Mexico); Camissonia (12; western North America to western South America), Eremothera (7; western North America, western Mexico), Neoholmgrenia (2; western North America), Camissoniopsis (14; southwestern United States, northwestern Mexico), Tetrapteron (2; western United States, northwestern Mexico). – Cosmopolitan, with their highest diversity in western United States and northwestern Mexico. Intraxylary phloem present. Hypanthium long, caducous. Flowers usually tetramerous (in Circaea dimerous). Sepals reflexed, caducous. Stamens rarely one or two. Transseptal vascular bundles present. Style sometimes short, often without vascular bundles. Stigma usually entire (sometimes quadrilobate, Dry type). Parietal tissue occasionally ten to c. 25 cell layers thick. Fruit sometimes a berry. n = 5–11, 15, 18?

Phylogeny of Onagraceae based on DNA sequence data and morphology (Levin & al. 2003, 2004; Wagner & Hoch 2005 onwards).

PENAEACEAE Sweet ex Guill.

( Back to Myrtales )

Guillemin in J. B. G. M. Bory de Saint-Vincent, Dict. Class. Hist. Nat. 13: 171. 1 Mar 1828, nom. cons.

Penaeales Lindl., Nix. Plant.: 15. 17 Sep 1833

Genera/species c 7/23

Distribution Southwestern South Africa, with their largest diversity in the southwestern parts of the Western Cape Province.

Fossils Unknown.

Habit Bisexual, small, evergreen, often procumbent shrubs with usually ericoid leaves (sometimes with lignotuber). Many species are xerophytes.

Vegetative anatomy Phellogen ab initio pericyclic. Cortical vascular bundles present or absent. Primary medullary rays narrow. Polyderm present. Endodermis sometimes with large cells. Vessel elements with simple perforation plates; lateral pits usually alternate (rarely opposite), simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements usually tracheids (rarely fibre tracheids) with bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular? Axial parenchyma paratracheal scanty or vasicentric, sometimes diffuse (apotracheal parenchyma absent). Intraxylary phloem present. Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate druses or crystal clusters often present.

Trichomes Hairs usually absent (unicellular simple hairs sometimes present).

Leaves Opposite, simple, entire, coriaceous, often ericoid, with ? ptyxis. Stipules axillary, rudimentary, colleter-like, or absent; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate, usually brochidodromous; midvein sometimes with apical swelling or glandular apex. Stomata anomocytic. Cuticular wax crystalloids?, sparse or absent. Mesophyll with fibres (sometimes with spiral thickenings) and sclerenchymatous idioblasts with calciumoxalate druses. Sclereids abundant. Leaf margin usually entire (in Sonderothamnus serrate).

Inflorescence Axillary, cymose or racemose, or flowers paired or solitary. Bracts often coloured, petaloid. Floral prophylls (bracteoles) in one or several pairs.

Flowers Actinomorphic. Hypanthium campanulate or cylindrical, persistent, often coloured. Half epigyny. Sepals four, with simple-valvate or reduplicate-valvate (valvate with recurved margins) aestivation, usually petaloid, persistent, free. Petals absent. Nectar secreted from glands along lower part of adaxial side of hypanthium. Disc absent.

Androecium Stamens four, obhaplostemonous, alternisepalous. Filaments short, adnate to hypanthial margin, sometimes inflexed in bud, free from each other and from tepals. Anthers basifixed, sometimes versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective laminar, distally (abaxially) expanded, often much longer than thecae; endothecium ephemeral. Tapetum secretory, with finally binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3–5(–10)-colporate, heterocolpate (with pseudocolpi or intercolpate pits alternating with colpi), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, rugulate or psilate (sometimes punctate).

Gynoecium Pistil composed of four connate antesepalous carpels. Ovary semi-inferior, quadrilocular. Style single, simple, subulate or quadrangular in cross-section (in Penaea and Stylapterus with four longitudinal commissural wings or ridges, respectively, running from ovary to stylar apex). Stigma usually capitate or quadrilobate, commissural (in Penaea and Stylapterus with four subapical stigmatic areas in angles of wings or ridges, respectively), Wet type? Pistillodium absent.

Ovules Placentation basal, axile or basal-apical. Ovules two to four (to numerous) per carpel, anatropous, ascending and/or pendulous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument two cell layers thick. Parietal tissue three or four cell layers thick. Megagametophyte tetrasporous, 16-cellular, Penaea type. Endosperm development ab initio nuclear. Chalazal endosperm haustorium present. Embryogenesis onagrad or asterad? (Penaea).

Fruit A loculicidal capsule, surrounded by persistent hypanthium.

Seeds Aril absent. Elaiosome formed by funicle. Testa two-layered. Exotesta usually well developed (sometimes crushed). Endotestal cells very prolonged; endotesta sometimes collapsed and represented by crystalliferous layer. Exotegmen and endotegmen fibrous or crushed. Perisperm not developed. Endosperm entirely absent or almost absent. Embryo straight, well differentiated, without suspensor, chlorophyll? Hypocotyl large. Cotyledons two, very small. Germination phanerocotylar?

Cytology n = 10, c. 20

DNA

Phytochemistry Very insufficiently known. Non-hydrolyzable tannins present. Aluminium not accumulated.

Use Ornamental plants, medicinal plants.

Systematics Endonema (2; southwestern Western Cape); Glischrocolla (1; G. formosa; southwestern Western Cape), Sonderothamnus (2; southwestern Western Cape), Saltera (1; S. sarcocolla; southwestern Western Cape), ’Brachysiphon’ (5; southwestern Western Cape; non-monophyletic), ’Stylapterus’ (8; southwestern Western Cape; non-monophyletic), Penaea (4; southwestern and southern Western Cape, southern Eastern Cape).

Penaeaceae are sister to Olinia (Oliniaceae).

Endonema is sister to the remaining Penaeaceae and the clade [Glischrocolla+Brachysiphon rupestris] is sister-group to the remainder. Sonderothamnus, Sarcocolla, and Brachysiphon mundii form a clade above these. Penaea appears to be monophyletic, although P. dahlgrenii is more closely allied to Stylapterus ericoides (Schönenberger & Conti 2003).

Cladogram of Penaeaceae based on DNA sequence data (Schönenberger & Conti 2003).

RHYNCHOCALYCACEAE L. A. S. Johnson et B. G. Briggs

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Johnson et Briggs in Ann. Missouri Bot. Gard. 71(1984): 732. 19 Apr 1985

Genera/species 1/1

Distribution Southeastern South Africa.

Fossils Unknown.

Habit Bisexual, evergreen trees. Young stems and branches terete or flattened in cross-section.

Vegetative anatomy Phellogen ab initio cortical. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits? Imperforate tracheary xylem elements libriform fibres usually with simple pits (almost without bordered pits), septate or non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma paratracheal scanty vasicentric (apotracheal parenchyma absent). Phloem with scattered sclereids. Sieve tube plastids Ss type? Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate druses present. Fibres with prismatic crystals.

Trichomes Hairs absent.

Leaves Opposite, simple, entire, coriaceous, with ? ptyxis. Stipules rudimentary, colleter-like, on petiole base, intrapetiolar (axillary) or absent; leaf sheath absent. Petiole vascular bundle transection arcuate; petiole with branched sclereids. Venation pinnate, eucamptodromous; midvein with apical gland. Stomata laterocytic, intermediate between anomocytic and cyclocytic. Cuticular wax crystalloids? Mesophyll without sclerenchymatous idioblasts. Leaf margin entire.

Inflorescence Usually terminal (sometimes axillary), panicle.

Flowers Actinomorphic, small. Hypanthium short. Hypogyny to almost half epigyny. Sepals (five or) six (or seven), with valvate aestivation, pointed, persistent, free. Petals (five or) six (or seven), with valvate? aestivation, alternisepalous, clawed, lobate, caducous, free. Nectary absent. Disc absent.

Androecium Stamens (five or) six (or seven), obhaplostemonous, alternisepalous, antepetalous. Filaments long, flattened, inserted immediately below petals on adaxial margin of hypanthium, inflexed in bud, free from each other and from tepals. Anthers subbasifixed (basifixed to dorsifixed), versatile, tetrasporangiate (microsporangia lateral), introrse, longicidal (dehiscing by longitudinal slits); loculi opening separately; septum between two microsporangia of each theca persistent also during anther dehiscence; connective little expanded abaxially; endothecium ephemeral. Tapetum secretory, with finally binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, heterocolpate (with pseudocolpi alternating with apertures), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, perforate or punctate.

Gynoecium Pistil composed of two (or three) connate carpels. Ovary superior to semi-inferior, bilocular (or trilocular), dorsiventrally compressed. Transseptal vascular bundles present. Style single, simple, short, stout, with persistent lower part. Stigma capitate to punctate, papillate, Wet type? Pistillodium absent.

Ovules Placentation parietal. Ovules c. 15 to c. 20 (to c. 50?) per carpel, anatropous, horizontal, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue three or four cell layers thick. Nucellar cap approx. three cell layers thick. Archespore multicellular. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit An apically loculicidal capsule, dorsiventrally compressed.

Seeds Aril absent. Seed coat testal. Testa winged at micropylar end, membranous; all testal cells persistent. Exotestal cells tanniniferous, with outer walls lignified. Endotesta? Tegmen? Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, flattened, chlorophyll? Cotyledons two, folded. Germination?

Cytology n = 10

DNA

Phytochemistry Very insufficiently known. Myricetin not found. Aluminium accumulated.

Use Unknown.

Systematics Rhynchocalyx (1; R. lawsonioides; sandstone areas in coastal regions of southern KwaZulu-Natal and Pondoland).

Rhynchocalyx is sister to [Oliniaceae+Penaeaceae].

VOCHYSIACEAE A. St.-Hil.

( Back to Myrtales )

Saint-Hilaire, Mém. Mus. Natl. Hist. Nat. 6, 4: 265. 1820 [’Vochisieae’], nom. cons.

Vochysiales Link, Handbuch 2: 60. 4-11 Jul 1829 [‘Vochysiaceae’]

Genera/species 7/200–210

Distribution Tropical West and Central Africa, Central and tropical South America.

Fossils Unknown.

Habit Bisexual, usually evergreen trees or shrubs (sometimes lianas, rarely herbs).

Vegetative anatomy Phellogen ab initio superficial or pericyclic. Primary stem with bicollateral vascular bundles. Medulla usually with sclerified vascular bundles (absent in Callisthene and Qualea). Pericyclic fibres few or absent. Secondary lateral growth normal or anomalous (via cylindrical cambium). Vessel elements with simple perforation plates; lateral pits alternate, usually bordered pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres usually with simple pits (in Vochysia sometimes bordered pits), septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal banded, or paratracheal scanty vasicentric, aliform, lozenge-aliform, winged-aliform, confluent, scalariform, reticulate, unilateral, or banded. Tyloses sometimes frequent. Premedullary diffuse interxylary phloem present in Erisma and Erismadelphus. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (in ‘Ruizterania’ clade of Qualea 3:3, trilacunar with three traces); leaf traces proceeding along stem before entering petiole. Medulla sometimes with secretory resinous canals. Sclereids and mucilage cells present. Silica bodies often abundant. Calciumoxalate crystals solitary prismatic, or in groups.

Trichomes Hairs usually unicellular, uniseriate, simple or furcate (rarely multicellular, stellate), often brown; glandular hairs?

Leaves Opposite or verticillate, simple, entire, coriaceous, with conduplicate ptyxis. Stipules small, cauline (in Qualea replaced by large colleter-like glands or associated with extrafloral nectaries); leaf sheath absent. Petiole vascular bundle transection? Venation pinnate, brochidodromous or eucamptodromous. Stomata anomocytic or paracytic. Cuticular wax crystalloids as groups of parallel platelets. Epidermis with or without mucilaginous idioblasts. Leaf margin entire.

Inflorescence Terminal or axillary, thyrse, panicle or raceme-like, composed of cincinni.

Flowers Transversely zygomorphic or asymmetrical (sometimes appearing actinomorphic). Pedicel in some species of Qualea with nectaries. Hypanthium absent. Epigyny (Erismeae) or secondary hypogyny (ovary primarily epigynous). Sepals five, usually with imbricate (sometimes cochlear) aestivation, unequal in size, persistent, connate at base, one adaxial-lateral larger sepal with nectariferous receptacular spur (three sepals in Korupodendron petaloid), sometimes coiled. Petals usually one or five (rarely three or absent), with contorted or imbricate aestivation, unequal in size, clawed, caducous, free (Callisthene and Qualea with displaced petal). Nectary present on sepal (see above). Disc absent.

Androecium Fertile stamen single, straight, antepetalous (opposite abaxial lateral petal; in Qualea and Callisthene offset and seemingly antesepalous), principally obhaplostemonous. Filament free from tepal. Anther dorsifixed or basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia (one or) two (in Erisma, most species of Vochysia and some species of Qualea; in Salvertia and some species of Vochysia four) or absent (in some species of Qualea).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely syncolporate), not heterocolpate (pseudocolpi absent), shed as monads, ?-cellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate or striate.

Gynoecium Pistil composed of usually three (rarely four) connate carpels; median carpel adaxial. Ovary inferior or secondarily superior (in mature flowers), unilocular (Erismeae; in Erisma unilocular; at least in Erismadelphus due to pseudomonomery: one or two carpels degenerating), or trilocular. Style single, simple, sunken. Stigma terminal or lateral, subcapitate to punctate, usually papillate (in Vochysia with multicellular uniseriate hairs), Wet type. Pistillodium absent.

Ovules Placentation axile or apical. Ovules one or two (Erismeae) to numerous per carpel, hemianatropous, pendulous, epitropous, bitegmic, crassinucellar. Micropyle bistomal (in Qualea exostomal?). Outer integument two or three cell layers thick. Inner integument approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule or one-seeded samaroid fruit (pseudosamara, Erismeae) with four (Erisma) or five persistent calyx lobes modified into wings.

Seeds Aril absent. Testa winged (wing in Qualea and Callisthene formed as extension of testa; unilateral wing in Vochysia and Salvertia formed from compressed hairs), thin, mesotesta sclerified?, endotestal cells thickened, with pectine, mesotegmic cells fibrous, often with thickened walls, or testa multiplicative, exotesta provided with thickened hairs. Some layers persistent. Remaining testal layers and tegmen unorganized. Qualea and Callisthene with endotesta containing crystalliferous layer, fibrous exotegmen and tanniniferous endotegmen. In Vochysia inner integument and inner crystalliferous layer of outer integument resorbed. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, plicate or planoconvex (Erismeae). Germination?

Cytology n = 11, 12

DNA

Phytochemistry 5-deoxyflavonoids, ellagic acid and methylated ellagic acids, tannins, and anthraquinones present. Aluminium accumulated in at least some species.

Use Timber, medicinal plants, gums, seed oils (Erisma).

Systematics Salvertieae Bill., Hist. Plant. 5: 100, 101. late 1873 vel prim. 1874. Salvertia (1; S. convallariodora; southern Brazil). – Vochysieae Dumort., Anal. Fam. Plant.: 41. 1829. Vochysia (105–110; Central America, tropical South America). – QualeeaeQualea (c 60; Central America, tropical South America), Callisthene (14; southern and central Brazil, northern Paraguay, eastern Bolivia). – Erismeae Dumort., Anal. Fam. Plant.: 41. 1829. Erisma (16–20; tropical South America), Erismadelphus (2; tropical West and Central Africa), Korupodendron (1; K. songweanum; Central Africa).

Vochysiaceae are sister to Myrtaceae.

There is no comprehensive phylogeny of Vochysiaceae available. Erismeae are probably monophyletic. The phellogen is subepidermal to cortical. The epigynous flower has a single carpel with one or two lateral to apical ovules. The fruit is a pseudosamara (samaroid) with persistent and accrescent calyx. The seed has an undifferentiated multilayered vascularized testa.


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