COMMELINANAE Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 514. 4 Feb 1967


[Commelinales+Cannales]


CANNALES R. Br. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 269. Jan-Apr 1820 [‘Canneae’]

Zingiberales Griseb., Grundr. Syst. Bot.: 167. 1-2 Jun 1854 [‘Zingiberides’]; Zingiberidae Cronquist in Brittonia 30: 505. 19 Dec 1978; Zingiberanae Takht. ex Reveal in Phytologia 74: 179. 25 Mar 1993

Fossils Spirematospermum is represented by leaves, trilocular fruits and seeds from the Campanian to the Maastrichtian and the Cenozoic of Europe and North America. The seeds are spirally striate, operculate and have a chalazal chamber and a micropylar collar. Several zingiberalean fossils have been described from the Maastrichtian Deccan Intertrappean Beds in India, such as Heliconiaites, Cyclanthodendron, Musocaulon and Heliconites, which are fossilized leaves and stems, and Musa cardiosperma, which comprises fruits with seeds, although it is questioned whether these should be attributed to Musa. Tricostatocarpon and Striatornata are trilocular fruits and operculate seeds from the Campanian of Mexico. Leaves from Late Cretaceous layers in the United States have been described as Zingiberopsis magnifolia, and some fossil seeds from the Eocene of Oregon resemble extant Ensete.

Habit Bisexual, perennial, often giant, herbs (stem rarely lignified), often with a pseudostem consisting of superimposed leaf sheaths. Sometimes with stilt roots. Rhizome and roots usually rich in starch.

Vegetative anatomy Roots usually with V-shaped aggregations of xylem, with large metaxylem element in angle. Phellogen absent. Stem cortex and central tissue with each vascular bundle surrounded by dense fibre envelope; cortex and central tissue separated by cylinder of fibres. Secondary lateral growth absent. Vessels present in roots (rarely also in stem). Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids usually P2cs type (with starch grains and cuneate protein crystals; sometimes P2cfs type, with starch grains, and cuneate protein crystals and filaments). Nodes multilacunar with several leaf traces. Stem sometimes with lysigenous mucilaginous cavities and ducts. Tanniniferous idioblasts often frequent. Idioblasts with suberised cell walls and containing aromatic (ethereal) oils and resins. Silica often present in vascular bundle sheaths. Silica bodies druse-like, stellate, conical or boat-shaped (sometimes present in stegmata), or absent. Calciumoxalate as acicular, rhomboidal, prismatic or tubular crystals, or as raphides or crystal sand. Starch grains usually simple (sometimes compound), often flat (sometimes irregular), usually excentric (sometimes compound; rarely spherical or elliptic).

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched (rarely furcate to stellate or candelabra-like), or absent.

Leaves Alternate (usually distichous, sometimes tetrastichous or spiromonostichous), simple, entire, often differentiated into pseudopetiole and pseudolamina, with supervolute ptyxis (laterally rolled-up in bud). Stipules absent; leaf sheath usually open (sometimes closed), with or without ligule at distal end. Pseudopetiole bundle transection arcuate. Venation pinnate-parallelodromous (with midvein, S-shaped lateral veins and fine transverse venation). Stomata usually tetracytic or paracytic, with symmetrical guard cells (rarely polycytic, hexacytic or tricytic); neighbouring cells with non-oblique divisions, or annular cells with non-oblique or oblique divisions. Cuticular wax crystalloids as platelets or longitudinally aggregated rodlets (Strelitzia type), or absent. Mesophyll with idioblasts containing ethereal oils and/or calciumoxalate as druses, prismatic or rhomboidal crystals or crystal sand (raphides usually absent). Epidermis cells often with spherical silica bodies or silica sand. Leaf margin entire.

Inflorescence Usually terminal (rarely axillary), capitate or cone-like, or simple or branched thyrse (sometimes raceme-like or spicate), consisting of few- to many-flowered (sometimes single-flowered) cymules/cincinni subtended by spiral bracts (bracts rarely absent). Partial inflorescences sometimes consisting of enantiomerous cincinni. Bracts persistent, often large. Floral prophylls (bracteoles) sometimes lateral, sometimes absent.

Flowers Zygomorphic or asymmetrical. Epigyny. Tepals 3+3; outer tepals with imbricate aestivation, usually sepaloid, free or more or less connate into tubular or spatha-like structure; inner tepals with imbricate aestivation, petaloid, more or less connate into tube; when zygomorphic, then median outer tepal abaxial. Gynoecial/septal nectaries epigynous, inserted at or sunken into ovary apex (sometimes labyrinthine); one or three glandular nectaries often present at stylar and staminal bases (sometimes also androecial nectaries on staminodia). Disc usually absent.

Androecium Stamen one, five, 3+2 or 2+3 (rarely 3+3) fertile, and one to four staminodial (rarely absent), often petaloid. Filaments free or connate at base, usually adnate to tepal base (sometimes free or absent). Anthers dorsifixed or basifixed, versatile or non-versatile, disporangiate or tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by pores). Tapetum secretory or amoeboid-periplasmodial. Staminodia one to five, variously arranged, often petaloid, free or more or less connate, sometimes fused with fertile stamen (sometimes absent).

Pollen grains Microsporogenesis usually successive (sometimes simultaneous). Pollen grains usually inaperturate (sometimes monosulcate, disulcate, sulcatoporate, 5–16-pantoporate, or spiraperturate), shed as monads, bicellular or tricellular (rarely unicellular) at dispersal. Exine tectate, with columellate infratectum, or intectate, consisting of scattered columellae or spinules, verrucate, gemmate, echinate, scabrate, rugulate, striate or smooth, or exine absent.

Gynoecium Pistil composed of usually three (rarely two) eusyncarpous or paracarpous connate carpels; median carpel abaxial. Ovary inferior, usually trilocular (later often incompletely or entirely trilocular or unilocular, rarely bilocular). Style single, simple, filiform, hollow, inserted in groove in filament of fertile stamen and between thecae. Stigma infundibuliform, cupular, decurrent or trilobate (rarely bilamellate), papillate or non-papillate, Wet type. Pistillodium absent.

Ovules Placentation axile, basal or subbasal (when ovary trilocular) or parietal (when ovary unilocular), rarely basal due to reduction of septa. Ovules four to more than 50 per carpel, anatropous or orthotropous (finally sometimes amphitropous), apotropous, bitegmic, crassinucellar or weakly crassinucellar. Micropyle usually endostomal (sometimes bistomal or exostomal). Cylindrical thickening, ’micropylar collar’, well developed? Parietal tissue one or two cell layers thick, formed from archesporial cell (sometimes absent). Nucellar cap often present. Epidermal cells of megasporangium often radially elongate and forming ‘nucellar pad’. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral, not proliferating. Endosperm development ab initio nuclear or helobial; chalazal chamber often early degenerating. Endosperm haustoria? Embryogenesis asterad, caryophyllad or chenopodiad.

Fruit Usually a loculicidal, dry or fleshy, capsule or a drupe (sometimes an irregularly dehiscing capsule, or a nut; rarely a berry or a syncarp with drupaceous or baccate mericarps).

Seeds Aril usually present. Seed coat endotestal. Exotesta consisting of fibriform elongate cells and often with phytomelan layer on epidermal cell walls (sometimes lignified or sclerenchymatous). Mesotesta sclerotic, tanniniferous or absent. Endotestal cells with silica, usually sclerotic, often with thickening U-shaped in cross-section, or absent. Operculum formed from inner testal epidermis, present near radicula and combined with ‘micropylar collar’ developed from outer integument and forming annular invagination in perisperm surrounding operculum. Tegmen thin, undifferentiated, often crushed. Perisperm well developed, rich in compound starch grains. Chalazosperm well developed or absent. Endosperm usually sparse, with aleurone (sometimes with lipids and/or starch grains). Embryo usually straight (sometimes curved or plug-shaped), often well differentiated, without chlorophyll or lipid bodies, Trillium type. Cotyledon one, ligulate. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode short to long. Mesocotyl absent. Coleoptile present or absent. Collar usually indistinct. Collar rhizoids sometimes present. Germination phanerocotylar? or cryptocotylar.

Cytology n = 7–14, 17, 21, 25 (x = 11)

DNA Plastid gene atpA with 6 bp deletion. Triplication of nuclear CYC-like gene. Duplications of GLOBOSA-like genes.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, isorhamnetin, syringetin), O-methylated flavonols, flavonol sulfates, dihydroflavones, flavone-C-glycosides, flavanones, luteolin, chalcones, cyanidin, delphinidin, pelargonidin, malvidin, etc., anthoxanthins, deoxyanthocyanins (apigenin etc.), ethereal oils consisting of mono-, di-, tri- and sesquiterpenes, proanthocyanidins (prodelphinidin, procyanidin), phenylpropanoids and related curcumins (diarylheptanoids), caffeic acid, chelidonic acid, zingerone (phenol), indole alkaloids, quinines, glycosides (e.g. 3-glycoside, 3-rutenoside), cyanogenic glycosides, steroidal saponins, resins, aromatic acids and ketones, benzoic quinones, shikimic acid- or polyacetate-derived arthroquinones, phenylphenalenones, and oxypipe colanic acid (polysaccharide). Cell walls with ferulic acid? Ellagic acid not found.

Systematics Zingiberales are sister-group to Commelinales with strong support.

The basal branching in Zingiberales is still unclear, although Musaceae are sometimes identified (with low support) as sister to the rest. Heliconia (Heliconiaceae) is part of the basal polytomy. Strelitziaceae and Lowiaceae appear to be sister-groups. According to Stevens (2001 onwards) these share the following potential synapomorphies: pseudopetiole with adaxial and abaxial series of air canals; perianth whorls coloured, distinct; adaxial stamens of inner whorl sterile; floral column (sterile apex of ovary) present; stigma trilobate; aril hairy; and exostomal aril lobed or fimbriate.

A well supported clade is [[Cannaceae+Marantaceae]+[Zingiberaceae+Costaceae]], which is characterized by the following features (Stevens 2001 onwards): silica bodies druse-shaped, boat-shaped; absence of raphides; pseudopetiole short and poorly differentiated, with a single series of air canals; stomatal guard cells with inner and outer ledges unequal (asymmetrical in transverse section); inner tepals connate; adaxial stamen of inner whorl fertile; tapetum amoeboid-periplasmodial, non-syncytial; both androecial whorls with two staminodia; micropyle endostomal; ‘micropylar collar’ well developed; exotestal cells longitudinally elongate; endosperm sparce; and presence of chalazosperm.

The [Cannaceae+Marantaceae] clade has the synapomorphies (Stevens 2001 onwards): pseudopetiole with oblique cells; flowers asymmetrical, pairwise; only one staminal half fertile; staminodia free; stigma not very expanded; endosperm absent or almost so; cells of chalazal intrusion into megasporangium degenerating and forming chalazal canal; and x=9.

The clade [Zingiberaceae+Costaceae] is characterized by: ligulate leaves; outer tepals connate; nectaries two, at apex of ovary; labellum formed by lateral staminodia of outer staminal whorl and by two staminodia of inner whorl; labellum large, with narrow tube and distinct open limb; exine present; style slender, running between two anthers halves; stigma cupular or infundibular; endosperm development helobial; chalazal mass in seed developed; endosperm persistent, not very abundant; and hypocotyl well developed.

Cladogram of Cannales based on DNA sequence data.

CANNACEAE Juss.

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de Jussieu, Gen. Plant.: 62. 4 Aug 1789 [’Cannae’], nom. cons.

Genera/species 1/8–10

Distribution Tropical America.

Fossils Unknown.

Habit Bisexual, perennial herbs, often giant herbs. Rhizome rich in starch, often tuberous.

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Stem in some species with numerous lysigenous mucilaginous cavities and canals. Tanniniferous idioblasts abundant. Silica bodies druse-shaped or stellate, adjacent to vascular bundles. Calciumoxalate as acicular, rod-shaped or plate-like or prismatic single crystals (raphides absent). Starch grains flat.

Trichomes Hairs absent.

Leaves Alternate (usually distichous, sometimes spiral), simple, entire, with supervolute ptyxis (laterally rolled-up in bud), often differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath open, without ligule. Venation pinnate-parallelodromous, with distinct mid-vein and numerous longitudinal primary veins. Stomata tetracytic, hexacytic or paracytic; guard cells with inner and outer ledges uniform. Cuticular wax crystalloids as longitudinally aggregated and often extremely long rodlets (Strelitzia type). Epidermis without silica bodies. Secretory cavities absent. Mesophyll without mucilaginous idioblasts; mesophyll cells with calciumoxalate as druses or single crystals (raphides absent). Leaf margin entire.

Inflorescence Terminal, simple or compound thyrse (sometimes raceme- or spike-like), usually with two-flowered cymules, cincinni; each cincinnus a pseudanthium with two asymmetrical flowers in each pair being mirrors of each other, enantiomery. Floral prophyll (bracteole) lateral, on right side of one flower and on left side of second flower in each pair.

Flowers Asymmetrical, large. Epigyny. Tepals 3+3; outer tepals sepaloid, persistent, free; inner tepals petaloid (one smaller), connate at base into a tubular structure. Septal/gynoecial nectaries sunken into ovary apex. Disc absent.

Androecium Stamen one fertile (adaxial outer stamen), petaloid, with one theca (two microsporangia), and (one to) three (or four) petaloid staminodia one inner of which, inserted opposite stamen, forming labellum and remaining ones forming ‘wings’; median adaxial inner stamen absent. Filaments somewhat connate at base, adnate to tepals. Anther dorsifixed, non-versatile, disporangiate (one theca degenerating), introrse, longicidal (dehiscing by longitudinal slits); placentoid present. Tapetum amoeboid-periplasmodial? or secretory?, with binucleate to sexanucleate cells. Staminodia usually three (rarely one, two or four).

Pollen grains Microsporogenesis simultaneous or successive. Pollen grains inaperturate, shed as monads, tricellular? or bicellular? at dispersal. Exine intectate, consisting of scattered columellae or spinules.

Gynoecium Pistil composed of three connate carpels; median carpel abaxial. Ovary inferior, trilocular, tuberculate to muricate. Style single, simple, flattened, petaloid, together with staminodia forming floral ‘upper lip’. Stigma terminal, decurrent along one stylar margin, papillate, Wet type. Pistillodium absent. Secondary pollen presentation (pollen grains deposited on abaxial side of style prior to anthesis).

Ovules Placentation axile. Ovules c. 20 to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument approx. ten cell layers thick. Inner integument ? cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ (‘Mikropylarkragen’), formed from outer integument surrounding micropyle. Hypostase present. Parietal cell? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit Usually a loculicidal capsule, tuberculate to muricate and glandular (sometimes a nut), with persistent outer tepals.

Seeds Seed pachychalazal (mitosis taking place in chalaza and basal part of megasporangium during ovule development). Aril probably as hairs from funicle; seed coat impermeable since exotesta and chalazal epidermis form a protecting layer of Malpighian-like cells; germination slit often developed adjacent to raphe, uplifted and hence permitting water uptake of seed. ‘Micropylar collar’ absent. Seed coat testal. Exotesta consisting of fibriform elongate Malpighian-like cells, without phytomelan layer. Mesotesta sclereidal. Endotesta and tegmen absent. Perisperm well developed, hard, copious, starchy. Chalazal tissue, chalazosperm, well developed. Endosperm one cell layer thick, with aleurone. Embryo straight, well differentiated, Trillium type, chlorophyll? Cotyledon one. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode short. Mesocotyl absent. Coleoptile present or absent (cotyledon sheath somewhat prolonged). Radicula well developed, persistent. Collar roots present. Germination?

Cytology n = 9 – Polyploidy occurring.

DNA

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, tannins, proanthocyanidins (prodelphinidin), chelidonic acid, and aromatic and other resins present. Ellagic acid, alkaloids, saponins, and cyanogenic compounds not found.

Use Ornamental plants, starch sources (South American or Queensland arrowroot from Canna edulis).

Systematics Canna (8–10; southeastern United States, Mexico, the West Indies, Central America, tropical and subtropical South America, with the largest diversity in northwestern South America).

The pollen grains in Canna are similar to those in its sister-group Marantaceae.

COSTACEAE (Meisn.) Nakai

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Nakai in J. Jap. Bot. 17: 203. Apr 1941

Genera/species 7/110–115

Distribution Pantropical, with their largest diversity in Central and South America, New Guinea and northern Australia.

Fossils Unknown.

Habit Bisexual, perennial (sometimes giant) herbs, often with aerial stem. Rhizome rich in starch. Root nodules rarely present. Rarely epiphytic. Rarely with axillary bulbils. Stem usually simple (in Tapeinochilos branched, with lateral branches penetrating leaf sheaths).

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vessels present in roots (sometimes also in stems) Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Ethereal oils absent. Raphide sacs and tanniniferous cells absent. Silica as stellate, druse-like or as single bodies. Calciumoxalate crystals frequent (raphides absent).

Trichomes Hairs unicellular or multicellular, uniseriate.

Leaves Alternate (tetrastichous or spiromonostichous: spirally arranged), simple, entire, with supervolute ptyxis (laterally rolled-up in bud). Stipules absent; leaf sheath closed, with a row of adaxial air canals (air canals absent from pseudopetiole and pseudolamina) and ligule in transition zone to pseudolamina. Vascular bundles adaxial. Hypodermis at least one-layered. Venation pinnate-parallelodromous; mid-vein distinct; secondary veins parallel, connected by transverse tertiary veins. Stomata tetracytic or paracytic; guard cells with inner and outer ledges non-uniform. Cuticular wax crystalloids absent. Epidermis without silica bodies. Mesophyll without mucilaginous idioblasts; mesophyll cells with calciumoxalate as druses or single prismatic crystals (raphides absent). Leaf margin entire.

Inflorescence Usually terminal (sometimes axillary), usually dense, cone- or spike-like (flowers in Monocostus solitary, axillary). Bracts imbricate, persistent, often with abaxial extrafloral nectaries at apex. Floral prophyll (bracteole) lateral, tubular or plicate.

Flowers Zygomorphic, large. Epigyny. Tepals 3+3; outer tepals with imbricate aestivation, sepaloid, persistent, connate into a tubular perianth (sometimes spatha-like); perianth tube on one side split to base; median outer tepal abaxial; inner tepals with imbricate aestivation, petaloid, longer than outer tepals, unequal in size, connate at base into a tube; median inner tepal larger, often bent upwards. Septal/gynoecial nectaries two, sunken into ovary apex. Disc often present.

Androecium Stamen single (probably median adaxial inner stamen), often petaloid. Filament flattened, free from tepals. Anther dorsifixed?, usually inserted in middle or lower part of filament (in Tapeinochilos at apex), non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective often apically widened (in Tapeinochilos pectinate). Placentoid present. Tapetum secretory, with binucleate cells. Staminodia five including abaxial stamen in outer whorl, petaloid, connate into a labellum inserted opposite stamen and usually much longer than perianth (labellum in Tapeinochilos strongly reduced); stamen and labellum connate at base into a tube.

Pollen grains Microsporogenesis successive. Pollen grains disulcate to sulcatoporate (foraminate; Dimerocostus, Monocostus), 5–16-pantoporate (foraminate; Costus), or spiraperturate (Tapeinochilos), shed as monads, bicellular at dispersal. Exine thick (resistant against acetolysis), consisting of rodlets packed into layers and covering interapertural areas.

Gynoecium Pistil composed of usually three (rarely two) connate carpels. Ovary inferior, usually trilocular (rarely unilocular or bilocular). Style single, simple, filiform, usually more or less adnate to filament of fertile stamen and inserted inside a groove between two anther halves (thecae); stylar canal present. Stigma usually cupular with fimbriate margins (sometimes bilamellate, consisting of two ciliate structures with a dorsally bilobate appendage), papillate, Wet type. Pistillodium absent.

Ovules Placentation usually axile (rarely parietal, then ovary unilocular). Ovules c. 15 to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument five or six cell layers thick. Inner integument ? cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ (‘Mikropylarkragen’), formed from outer integument surrounding micropyle. Hypostase present. Epidermal cells of megasporangium vertically elongate. Parietal cell? Parietal tissue? Megagametophyte monosporous, Polygonum type (or tetrasporous, Adoxa type?). Endosperm development helobial. Endosperm haustoria? Embryogenesis caryophyllad.

Fruit Loculicidal or irregularly (rarely explosively) dehiscing capsule with persistent outer tepals (rarely nutlike).

Seeds Aril often present. Seed coat testal. Exotesta consisting of fibriform elongate cells? and with phytomelan layer on epidermal cell walls. Endotesta well developed, with cells rich in silica. Tegmen degenerated. Operculum present adjacent to radicula and combined with ‘micropylar collar’. Perisperm well developed, rich in starch (with single, concentric starch grains). Chalazal tissue, chalazosperm, well developed. Endosperm sparse, with oily lipids, without starch. Embryo large, straight, with lipid bodies, Trillium type, with chlorophyll. Cotyledon one, differentiated into one sheath-like and one lamina-like part, photosynthesizing, with apical outgrowth directed backwards. Cotyledon hyperphyll compact (small), not assimilating (present at apex of large foliaceous assimilating sheath). Hypocotyl internode long. Coleoptile absent. Collar? Germination phanerocotylar?

Cytology n = 9, 18 (Costus); n = 14 (Dimerocostus)

DNA

Phytochemistry Flavone-C-glycosides, steroidal saponins, cyanogenic compounds, and benzoquinones present.

Use Ornamental plants.

Systematics Chamaecostus (8; tropical South America), Dimerocostus (2; tropical America), Monocostus (1; M. uniflorus; Amazonian Peru), Paracostus (2; tropical West and Central Africal, Malesia), Hellenia (>4; tropical Asia), Tapeinochilos (16–20; the Moluccas, New Guinea to tropical Australia), Costus (c 75; tropical regions on both hemispheres).

Costaceae are sister-group to Zingiberaceae.

Cladogram of Costaceae based on morphology and DNA sequence data (Specht 2006).

HELICONIACEAE Vines

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Vines, Stud. Text-book Bot. 2: 561, 562. Mar 1895 [’Heliconieae’]

Genera/species 1/c 200?

Distribution Tropical America, fewer species in East Malesia, Melanesia and eastwards to Samoa.

Fossils Unknown.

Habit Bisexual, perennial (often giant) herbs. Rhizome starchy.

Vegetative anatomy Rhizome with endodermis-like layer. Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Laticifers absent. Silica bodies boat-shaped, in idioblasts, stegmata, associated with vascular bundles. Idioblasts, raphide sacs, with calciumoxalate raphides abundant. Crystal sand present or absent. Rhizome with cylindrical or ellipsoidal starch grains.

Trichomes Hairs multicellular, uniseriate or branched (sometimes candelabra-shaped).

Leaves Alternate (distichous), simple, entire, differentiated in pseudopetiole and pseudolamina, with supervolute ptyxis (laterally rolled-up in bud). Stipules absent; leaf sheath well developed, without distinct ligule. Venation pinnate-parallelodromous, with distinct midvein and arcuate, densely spaced secondary veins, converging at apex and connected to transversal tertiary veins. Stomata usually polycytic (sometimes paracytic or tetracytic); adjacent cells with oblique divisions. Cuticular wax crystalloids as longitudinally aggregated rodlets (Strelitzia type), chemically dominated by wax esters. Epidermis without silica bodies. Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides (and/or crystal sand). Leaf margin entire.

Inflorescence Terminal, spike-like thyrse consisting of few- to many-flowered cincinni, each subtended and often enclosed by large boat-shaped spathe-like green to brightly coloured and usually persistent bract. Floral bracts small and usually membranous. Floral prophyll (bracteole) keeled, lateral.

Flowers Obliquely zygomorphic (asymmetric). Epigyny. Tepals 3+3, petaloid, five tepals connate into a tubular upper lip; median outer tepal adaxial, fused with remaining tepals only at base. Septal nectaries, a triradiate apically closed crater opening by three pores at stylar base, present in axile tissue from below locules and upwards. Disc absent.

Androecium Stamens 2+3 fertile. Filaments filiform, free, adnate at base to perianth tube. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial, non-syncytial. Median outer stamen modified into subulate to petaloid staminodium, inserted in perianth tube opposite median outer tepal.

Pollen grains Microsporogenesis successive. Pollen grains inaperturate (functionally monoaperturate), heteropolar, shed as monads, tricellular at dispersal. Exine thin and reduced, absent or discontinuous at distal pole. Pollen surface spinulate to psilate.

Gynoecium Pistil composed of three connate carpels. Ovary inferior, trilocular. Style single, simple, filiform, hollow, often somewhat curved. Stigma one, capitate or trilobate, papillate, Wet type. Pistillodium absent.

Ovules Placentation subbasal-axile. Ovule one per carpel, anatropous, apotropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument six to nine cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ (‘Mikropylarkragen’), formed from outer integument, surrounds micropyle. Inner integument two cell layers thick. Hypostase absent. Parietal cell formed from archesporial cell, not forming a distinct parietal layer or parietal tissue early degenerating. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Usually a drupe with one pyrene per locule (sometimes a schizocarp with two or three drupaceous mericarps; sometimes on prolonged pedicel). Mechanical layer mainly formed by endocarp.

Seeds Aril absent or almost so. Micropylar operculum formed from funicle and combined with ‘micropylar collar’. Testa thin, undifferentiated. Tegmen thin, undifferentiated. Perisperm one-layered, well developed, with starch and aleurone, surrounding endosperm. Endosperm copious, starchy and oily (with single, concentric starch grains). Embryo straight or slightly curved, little differentiated, Trillium type, chlorophyll? Cotyledon one. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode short. Coleoptile absent; leaf sheath lobed. Collar at right angles to cotyledon. Germination?

Cytology n = (11) 12

DNA

Phytochemistry Insufficiently known. Tannins and proanthocyanidins present. Cyanogenic compounds not found.

Use Ornamental plants.

Systematics Heliconia (100–200; Sulawesi and the Moluccas in East Malesia, Melanesia eastwards to Fiji, Samoa, Mexico, Central and South America, with their highest diversity in tropical America).

The species delimitations in Heliconia are notoriously difficult.

The abaxial outer stamen is sterile, a feature similar to that in the [[Cannaceae+Marantaceae]+ [Zingiberaceae+Costaceae]] clade. Heliconia is sister-group to the remaining Zingiberales in the study by Janssen & Bremer (2004), but the support for this placement is generally low.

LOWIACEAE Ridl.

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Ridley, Fl. Malay Penins. 4: 291. 1 Dec 1924, nom. cons.

Lowiales Takht. ex Reveal et Doweld in Novon 9: 551. 30 Dec 1999

Genera/species 1/16

Distribution Southernmost China, Indochina, West and North Malesia, West Pacific islands.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome rich in starch.

Vegetative anatomy Rhizome with endoderm-like layer on inner side of cortex; outer cortex often with fibre strands. Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Idioblasts, stegmata, with conical silica bodies in association with vascular bundles in stem and leaves. Calciumoxalate as raphides and single crystals. Rhizome cells with crystal sand. Starch grains compound.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, differentiated into pseudopetiole and pseudolamina, with supervolute ptyxis (laterally rolled-up in bud). Stipules absent; leaf sheath well developed, without ligule. Venation pinnate-parallelodromous, with distinct midvein and parallel secondary veins, connected with transverse tertiary veins; longitudinal veins abruptly ascending and converging towards apex. Stomata paracytic; guard cells deeply sunken, asymmetrical (with inner and outer borders unequal). Cuticular wax crystalloids absent. Epidermis without silica bodies. Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, cymose, consisting of repeated one- or few-flowered partial inflorescences, branching from scale-like bracts below flower. Prophyll two-keeled; above these two interphyll with axillary partial inflorescence buds; above these a bract with an axillary flower; on top a reduced bract enclosing inflorescence apex.

Flowers Zygomorphic, often resupinate and then with median inner tepal adaxial, more or less evil-smelling. Epigyny. Tepals 3+3; outer tepals sepaloid, connate at base into a tubular structure; median outer tepal adaxial; inner tepals petaloid, free; two lateral inner tepals smaller and sometimes narrow and enclosing stamens; median inner tepal modified into large sometimes boat-shaped adaxial labellum (in resupinate flowers abaxial). Septal/gynoecial nectaries sunken into upper part of ovary. Disc absent.

Androecium Stamens five (median adaxial absent), in a single whorl. Filaments free, adnate at base to inner tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective somewhat prolonged at apex. Tapetum secretory? Staminodium usually absent.

Pollen grains Microsporogenesis successive? Pollen grains inaperturate, shed as monads, ?-cellular at dispersal. Exine intectate, psilate.

Gynoecium Pistil composed of three connate carpels. Ovary inferior, trilocular, prolonged at apex. Style single, simple, elongate, widened at apex. Stigma trilobate, zygomorphic, dorsiventrally flattened, with secretory tissue, ‘viscidium’, at base of adaxial/ventral side and with entire, lobate or fimbriate stigmatic lobes, non-papillate, Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules c. 30 to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument 14 to 16 cell layers thick. Inner integument approx. four cell layers thick. ‘Micropylar collar’ developed? Parietal cell? Megagametophyte monosporous, Polygonum type? Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule with lower part of ovary persistent.

Seeds Aril lacerate, consisting of three or four coarse hair-like outgrowths. Testa vascularized, with operculum opposite radicula. Seed coat testal. Exotesta and adjacent two inner layers lignified; exotesta with phytomelan on cell walls. Endotesta silicified?, consisting of radially elongate sclereids. Tegmen crushed? Perisperm thin, slightly developed. Endosperm copious, with starch grains. Embryo cylindrical, well differentiated, Trillium type, chlorophyll? Cotyledon one, not assimilating? Cotyledon hyperphyll? Hypocotyl internode? Coleoptile? Collar? Germination?

Cytology n = 9

DNA

Phytochemistry Virtually unknown. Tannins present. Cyanogenic compounds not found.

Use Unknown.

Systematics Orchidantha (16; southernmost China, Hainan, Indochina, Malaysia, Borneo, the Philippines, West Pacific islands).

Orchidantha is probably sister to Strelitziaceae (Cron & al. 2012).

MARANTACEAE R. Br.

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Brown in M. Flinders, Voy. Terra Austral. 2: 575. 19 Jul 1814 [’Maranteae, or Canneae’], nom. cons.

Marantales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 10. Sep-Oct 1835 [‘Marantaceae’]

Genera/species 25–30/520–530

Distribution Pantropical except Australia, with their largest diversity in tropical America.

Fossils Unknown.

Habit Bisexual, perennial (sometimes giant) herbs (rarely climbing; sometimes with woody rhizome). Rhizome rich in starch.

Vegetative anatomy Roots usually fibrous (rarely with nodules). Phellogen absent. Secondary lateral growth absent. Vessels present in roots (sometimes also in stems). Vessel elements with scalariform and/or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays usually absent. Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Mucilage cells and secretory cells rare. Mucilage canals usually absent (present in Thalia). Silica bodies with irregular outgrowths, conical or druse-like, in stegmata or thin-walled cells. Calciumoxalate as single, rhomboidal, prismatic or rod-shaped crystals (raphides absent). Starch grains usually elliptic and somewhat excentric (rarely flat or angular), single or aggregated.

Trichomes Hairs usually absent (sometimes unicellular).

Leaves Alternate (distichous, sometimes in right angles to stem), simple, entire, with supervolute ptyxis (laterally rolled-up in bud?), usually differentiated into often long pseudopetiole (often absent) and asymmetrical pseudolamina, with a pulvinus (with oblique cells) in transition zone between pseudopetiole and pseudolamina. Stipules absent; leaf sheath closed, without ligule. Venation pinnate-parallelodromous, with distinct mid-vein and closely spaced parallel arcuate lateral veins converging and fused at leaf margin and connected by transverse tertiary veins. Stomata tetracytic or paracytic (sometimes tricytic, anomocytic or polycytic); guard cells with inner and outer borders non-uniform. Cuticular wax crystalloids usually absent (sometimes non-orientated). Epidermis without silica bodies. Mesophyll without mucilaginous idioblasts; mesophyll cells with calciumoxalate as druses or single prismatic crystals (raphides absent). Leaf margin entire. Extrafloral nectaries present at apex of leaf sheath in some species of Ctenanthe and Stromanthe.

Inflorescence Usually terminal (sometimes axillary), simple or branched; each partial inflorescence – usually a spike- or head-like thyrse – consisting of groups (subtended by bracts) of one or several (rarely numerous) usually two-flowered (in Monotagma and Monophrynium one-flowered) cymules; each cymule with a basal adaxial prophyll (sometimes also with a scale-like abaxial interphyll and often one or two adaxial bracteoles); each cymule an enantiomerous pseudanthium with two asymmetrical flowers in each pair being mirror images of each other. Floral prophyll (bracteole) absent or lateral: on right side of first flower and on left side of second flower in each pair, sometimes with apical extrafloral nectaries.

Flowers Asymmetrical, often small, resupinate. Epigyny. Tepals 3+3; outer tepals sepaloid, free; inner tepals petaloid, connate at base into a tube, adnate to androecium and style; median outer perianth member adaxial; median inner tepal often dome- or hood-shaped, and larger than the rest. Septal/gynoecial nectaries sunken at ovary apex. Disc absent.

Androecium Outer staminal whorl consisting of one or two lateral sometimes petaloid staminodia, or absent; median adaxial outer stamen absent. Inner staminal whorl consisting of one fertile (adaxial inner) stamen, with a single lateral theca (two microsporangia) and often with a lateral petaloid outgrowth (transformed sterile theca?), and two petaloid staminodia: one sensitive dome- or hood-shaped staminodium, staminodium cucullatum (usually lobed and with lateral appendages), enclosing style and stigma prior to pollination, and a second staminodium, staminodium callosum, (hard, fleshy and warty on adaxial side). Filaments of outer staminodia connate into a tube, adnate at base to tepals and style; filament of inner stamen and staminodia connate at base into a tube. Anther dorsifixed?, non-versatile, disporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial? (or secretory?), with ?-nucleate cells.

Pollen grains Microsporogenesis successive. Pollen grains inaperturate, shed as monads, bicellular at dispersal. Exine very thin or absent; pollen wall consisting of intine. Pollen surface psilate.

Gynoecium Pistil composed of three connate carpels, two of which often as sterile pistillodia. Ovary inferior, trilocular, with two locules often compressed and sterile. Style single, subulate, adnate at base to perianth-staminal tube; a flat adaxial subapical surface delimited on abaxial side by gland-like secretory swelling; style held under tension by staminodium cucullatum prior to pollination. Pollen receptive surface (‘stigma’) present on inner side of an infundibuliform subapical cavity, non-papillate, Wet type. Pistillodia present or absent. Secondary pollen presentation (pollen grains deposited from developing theca on adaxial secretory surface prior to anthesis).

Ovules Placentation basal to axile. Ovule one per carpel, ab initio anatropous, subsequently amphitropous (anacampylotropous), bitegmic, crassinucellar. Micropyle usually endostomal (in Phrynium bistomal). Outer integument six to eight (to twelve) cell layers thick. Inner integument ? (one or two?) cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ (‘Mikropylarkragen’), formed from outer integument surrounding micropyle and penetrating micropylar part of megasporangium. Operculum formed by periclinal cell divisions outside of ‘collar’. Parietal cell formed from archesporial cell. Chalazal (vascular) tissue growing into megasporangium. Nucellar cap (sometimes approx. two cell layer thick) formed from palisade-like megasporangial epidermis. Lateral epidermal cells dividing periclinally. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes proliferating. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis chenopodiad.

Fruit Usually a loculicidal capsule (in Sarcophrynium and Thaumatococcus baccate; in Thalia nutlike).

Seeds Aril formed from outer epidermis of outer integument near micropyle, or absent. Seed coat testal. Exotesta consisting of fibriform elongate cells? and with phytomelan layer on epidermal cell walls. Mesotesta tanniniferous. Endotestal cells with silica. Operculum formed from endotestal epidermis and combined with ‘micropylar collar’ (cf. Canna). Tegmen crushed, sometimes with thin elongate sclereids. Perisperm hard, copious, with thick-walled cells containing starch grains. Chalazal tissue, chalazosperm, often well developed, with a branched or simple perisperm canal formed in megasporangium by degradation of chalazal tissue. Endosperm rudimentary or absent. Embryo long, curved, without chlorophyll, Trillium type. Cotyledon one. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode absent. Mesocotyl sometimes present; collar inserted at right angles to cotyledon. Coleoptile present. Germination?

Cytology n = 9–13 (chromosome numbers uncertain!)

DNA Mitochondrial coxI intron present in several genera.

Phytochemistry Flavonols (kaempferol, quercetin), flavonol sulphates, flavone-C-glycosides, luteolin, anthocyanins, deoxyanthocyanins (apigenin), cyanidin, delphinidin, pelargonidin, tannins, and cyanogenic compounds present. Thaumatins (high-molecular proteins) present in aril (1.600 times sweeter than saccharose). Ellagic acid, chelidonic acid, and saponins not found.

Use Ornamental plants, starch sources (West Indian arrowroot from Maranta arundinacea), vegetables and thatching (Calathea), basketry (Ischnosiphon).

Systematics Sarcophrynium (3; tropical Africa), Megaphrynium (4; tropical Africa), Thaumatococcus (1; T. daniellii; tropical West Africa), Hypselodelphys (5; tropical Africa), Trachyphrynium (1; T. braunianum; tropical West Africa); Haumania (2; tropical Africa); Monotagma (37; tropical South America), Goeppertia (c 15?; tropical South America), Pleiostachya (2; Central America, Ecuador), Ischnosiphon (c 35; tropical America),’Calathea’ pro parte (c 285; tropical America); Thalia (5–7; tropical America), Donax (3–4; Southeast Asia, Malesia and eastwards to Vanuatu), Schumannianthus dichotomus (India to Southeast Asia and the Philippines), ’Phrynium’ (c 20; tropical Asia; paraphyletic); Halopegia (2; Congo, Southeast Asia, Java), Schumannianthus virgatus (southern India, Sri Lanka), Hylaeanthe (5–6; tropical South America), Maranta (c 30; tropical America), Myrosma (1; M. cannifolia; the West Indies, tropical South America), Saranthe (5–10; Brazil), Ctenanthe (c 10; Costa Rica, Brazil), Stromanthe (c 13; tropical South America); Ataenidia (1; A. conferta; tropical Africa), Marantochloa (c 15; tropical Africa, Madagascar, the Comoro Islands), Stachyphrynium (c 10; tropical Asia; incl. Afrocalathea?), Afrocalathea (1; A. rhizantha; tropical West Africa; in Stachyphrynium?); unplaced Monophrynium (3; Central Malesia), Monophyllanthe (1–2; northern South America), Cominsia (4; East Malesia, New Guinea, Vanuatu).

Marantaceae are sister-group to Canna (Cannaceae). The Sarcophrynium clade is sister to the remaining Marantaceae (Prince & Kress 2006).

The trend towards secondary pollen presentation may be a synapomorphy of Marantaceae and Canna. In Marantaceae it is a type of explosive mechanism. The pollen grains are deposited from the developing theca into a cavity on the adaxial side near the apex of the style, which is curved and held under tension by the staminodium cucullatum. The secretions from the adjacent glands make the pollen grains stick to the stylar surface. The appendages of the staminodium are touched by the nectar-searching pollinator, and the style triggered and momentarily (within a few thousands of a second) recurved releasing the deposited pollen. Simultaneously, pollen grains from a different flower may be transferred from the pollinator to the receptive surface.

Cladogram of Marantaceae based on DNA sequence data and morphology (Andersson & Chase 2001; Borchsenius & al. 2012).

Cladogram (one of 48 shortest maximum parsimony trees) of Marantaceae based on DNA sequence data (Prince & Kress 2006).

MUSACEAE Juss.

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de Jussieu, Gen. Plant.: 61. 4 Aug 1789 [’Musae’], nom. cons.

Musales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 269. Jan-Apr 1820 [‘Musae’]

Genera/species 2/40–45

Distribution Tropical Africa, Madagascar, tropical Asia, Queensland, West Pacific islands.

Fossils Uncertain. Fossil remains from the Eocene of Oregon have been assigned to Ensete.

Habit Usually monoecious or andromonoecious, with male flowers at apex and usually female flowers (rarely bisexual flowers) at base of inflorescence, perennial giant herbs with aerial pseudostem consisting of leaf sheaths (species in Ensete hapaxanthic, monocarpous). Rhizome rich in starch, with short and thick branches. Axillary buds leaf-opposite or absent.

Vegetative anatomy Root cortex sometimes with periderm. Root medulla with scattered wide vessels and phloem strands. Endodermis present in rhizome. Phellogen absent. Secondary lateral growth absent. Vessels present in roots and rhizome. Vessel elements with scalariform and/or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2cfs type, with cuneate protein crystals, starch grains and protein filaments. Articulated laticifers with mucilage (becoming dark when oxidized) numerous in Musa in association with vascular bundles (not in roots). Silica bodies boat-shaped, in association with vascular bundles (not so in roots). Idioblasts (raphide sacs) with calciumoxalate raphides. Starch grains simple, concentric.

Trichomes Hairs absent from vegetative parts.

Leaves Alternate (spiral), simple, entire, with supervolute ptyxis (laterally rolled-up in bud?), usually differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath well developed, without distinct ligule. Pseudopetiole with a single row of abaxial air canals. Venation pinnate-parallelodromous; mid-vein distinct; secondary veins closely spaced, parallel, arising in almost right angles from mid-vein, converging at leaf margin and connected by transversal tertiary veins. Stomata usually paracytic (sometimes tetracytic). Cuticular wax crystalloids as longitudinally aggregated rodlets (Strelitzia type), chemically dominated by wax esters or alkanes. Epidermis without silica bodies. Mesophyll with mucilaginous idioblasts containing calciumoxalate as raphides and single rhomboidal crystals. Leaf margin entire.

Inflorescence Terminal, compound spike-like thyrse consisting of lateral cincinni in axils of large, caducous (in Musa) or persistent (in Ensete), spatha-like and boat-shaped bracts. Thyrse with basal female (rarely bisexual) flowers and apical male flowers. Prophylls lateral. Floral bracts and prophyll (bracteole) absent. Extrafloral nectaries sometimes present.

Flowers Zygomorphic, large. Epigyny. Tepals 3+3, petaloid, pseudomonocyclic; median outer tepal abaxial (anterior); three outer and two lateral inner tepals connate into a tube or spatha; median inner tepal adaxial (posterior), free, small and dome-shaped (sometimes deeply concave), directed downwards. Septal/gynoecial nectaries present in upper part of ovary. Disc absent.

Androecium Stamens five, with inner median adaxial (posterior) stamen small or staminodial (in Musa), or stamens six (in Ensete), antetepalous. Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective occasionally somewhat prolonged. Tapetum secretory, with binucleate cells. Staminodium one (opposite free stamen) or absent; female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains inaperturate (omniaperturate), shed as monads, bicellular at dispersal. Exine very thin to almost absent; pollen surface psilate (in Musa) or gemmate to verrucate (in Ensete). Intine thick.

Gynoecium Pistil composed of three connate carpels; median carpel abaxial (anterior). Ovary inferior, trilocular, in Musa with mucilage and hairs on placentae (sometimes also on funicle). Style single, simple, filiform, often widened at apex. Stigma capitate, papillate, Wet type. Male flowers with pistillodium.

Ovules Placentation axile. Ovules approx. ten to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle sometimes exostomal. Outer integument six to nine cell layers thick. Inner integument two or three cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ or ‘Mikropylarkragen’, formed from outer integument surrounding micropyle. Hypostase present. Parietal cell not formed (parietal tissue absent). Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A berry-like capsule with leathery pericarp, indehiscent or partially dehiscing longitudinally. Seeds surrounded by a fleshy starchy pulp formed from placental hairs.

Seeds Funicular aril reduced, consisting of simple hairs embedded in mucilage, or absent. Seed with chalazal space. Exotestal cells silicified, forming dense sclerenchyma. Mesotesta massive, 20–25 cell layers thick, sclerotic. Endotestal cells silicified? Tegmen two cell layers thick, with elongate cells. Operculum situated above micropylar part and combined with ‘micropylar collar’. Perisperm one cell layer thick, with lipids (starch absent). Endosperm copious, starchy (with small chalazal space?). Embryo straight (in Musa) or somewhat curved (in Ensete), fungiform, Trillium type, chlorophyll? Cotyledon one. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode relatively well developed. Mesocotyl absent. Coleoptile present. Double micropylar collar well developed (at right angles to cotyledon). Germination phanerocotylar.

Cytology n = 7, 9–11 (Musa); n = 9, 10 (Ensete)

DNA Mitochondrial coxI intron present.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, pelargonidin, malvidin and malvidin glycoside, anthoxanthins, deoxyanthocyanins, tannins, proanthocyanidins (prodelphinidins), indole alkaloids, cyanogenic glycosides, and phenylphenalenones (perinaphthenones) present. Ellagic acid and saponins not found.

Use Ornamental plants, fruits and vegetables, textiles, carpets and rope (abaca, manila-hemp from Musa textilis), beer, etc.

Systematics Musa (35–40; eastern and southern India, Sri Lanka, Southeast Asia including southernmost China, Malesia, northeastern Queensland, islands in the western Pacific), Ensete (6; tropical and subtropical regions in Africa, Madagascar and Asia eastwards to New Guinea; incl. Musella).

Possibly, Musaceae are sister to the remaining Cannales, but the support is low.

Musella is nested within Ensete in analyses by Liu & al. (2010).

STRELITZIACEAE (K. Schum.) Hutch.

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Hutchinson, Fam. Fl. Pl. 2: 72. 20 Jul 1934, nom. cons.

Genera/species 3/6–7

Distribution South Africa, Madagascar, tropical South America.

Fossils Uncertain. The fossil genus Cyclanthodendron (including fruits with the name Tricoccites) from the Eocene of India? has sometimes been attributed to Cyclanthaceae, but possibly is more allied to Strelitziaceae.

Habit Bisexual, perennial (often giant) herbs (aerial stem in Phenakospermum, Ravenala and Strelitzia nicolai lignified). Rhizome rich in starch, with short and thick branches. Dichotomous branching present in Strelitzia.

Vegetative anatomy Root medulla with uniformly scattered wide xylem and phloem strands. Aerial stems with superficial mechanical layer and with vascular bundles surrounded by dense fibre envelopes. Phellogen absent? Central vascular cylinder not delimited by endodermis. Secondary lateral growth absent. Vessels present in roots and stem (in Ravenala only in roots). Vessel elements usually with scalariform (in Phenakospermum also simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Tannin frequent in normal cells. Silica bodies, druse-shaped or spherical, present in hypodermal cells in association with vascular bundles (not in roots), or spherical in epidermal stegmata. Idioblasts (raphide sacs) with calciumoxalate raphides. Starch grains simple, usually spherical (in Ravenala flattened).

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, with supervolute ptyxis (laterally rolled-up in bud?), usually differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath well developed, without distinct ligule. Pseudopetiole long, with several arcs of air canals. Venation pinnate-parallelodromous; midvein distinct; secondary veins closely spaced, parallel, converging at leaf margin and connected by transversal tertiary veins. Stomata polycytic, tetracytic or hexacytic, with sometimes oblique cell divisions and often sunken guard cells. Cuticular wax crystalloids as longitudinally aggregated terete rodlets (Strelitzia type), chemically dominated by wax esters, usually concentrated around stomata. Epidermis without silica bodies. Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal (Phenakospermum) or lateral (Ravenala, Strelitzia), compound spike-like thyrse consisting of one or several distichous cincinni in axils of large stout persistent spatha-like bracts. Bracts and floral prophylls (bracteoles) boat-shaped to lanceolate.

Flowers Zygomorphic, large. Epigyny. Tepals 3+3; median outer adaxial (posterior); two lateral inner tepals larger than median adaxial (cucullate) inner tepal; outer tepals free; inner tepals connate at base; in Strelitzia two lateral abaxial inner tepals connate into a sagittate structure surrounding stamens and style. Septal/gynoecial nectaries deeply sunken into upper part of ovary. Disc absent.

Androecium Stamens 3+2 (median inner stamen staminodial; Phenakospermum, Strelitzia) or 3+3 (Ravenala), antetepalous (at least in Strelitzia). Filaments free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective in Strelitzia somewhat prolonged; anthers in Strelitzia with filiform structures inside; anthers in Phenakospermum up to 7 cm long, present in perianth keel and first exposed at pollination (by bats). Tapetum secretory, with up to 32-ploid cells. Staminodium one (adjacent to adaxial inner tepal) or absent.

Pollen grains Microsporogenesis successive. Pollen grains inaperturate (omniaperturate), shed as monads, unicellular at dispersal (vegetative cell degenerating). Exine very thin to almost absent; pollen surface psilate? Intine thick. Pollen wall in Phenakospermum discontinuous.

Gynoecium Pistil formed by three connate carpels. Ovary inferior, trilocular, prolonged at apex. Style single, simple, filiform, often widened at apex. Stigma long, turbinate, or trilobate, papillate, Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules c. 20 to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Cylindrical ridge-shaped thickening, ’micropylar collar’ (‘Mikropylarkragen’), absent. Hypostase absent. Parietal cell absent or formed from archesporial cell, not forming distinct parietal layer. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A lignified loculicidal capsule.

Seeds Aril exostomal, fimbriate to lobate, sometimes brightly coloured, formed partly from outer integument near micropyle, partly from funicle opposite micropyle. Seed coat testal. Testa with compact outer cuticle. Inner testal epidermis consisting of large cells with inner tangential walls strongly thickened (U-cell layer). Tegmen consisting of cuticle (remaining layers crushed). Operculum rudimentary or absent. Perisperm one-layered, reduced, with lipids (starch absent), or absent. Endosperm copious, starchy, outer cell layers with aleurone. Embryo straight or curved, well differentiated, Trillium type, chlorophyll? Cotyledon one, massive. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode and mesocotyl absent. Coleoptile present. Collar? Radicula well developed, persistent. Germination?

Cytology n = 7, 9, 11

DNA

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, tannins, phenylphenalenones, and oxypipe colanic acid (in Strelitzia) present. Flavonols (kaempferol, quercetin) and cyanidin absent from Ravenala. Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Ravenala (1; R. madagascariensis; Madagascar), Phenakospermum (1; P. guyanense; eastern tropical South America), Strelitzia (4–5; temperate and subtropical regions in South Africa).

Strelitziaceae are probably sister to Orchidantha(Lowiaceae) (Cron & al. 2012).

Prince (2010) presents a cladogram of Cannales, including the following topology of Strelitziaceae: [Ravenala+[Phenakospermum+Strelitzia]]. On the other hand, Cron & al. (2012) were not able to unequivocally define the sister-group relationship in Strelitziaceae. Ravenala was sister to [Phenakospermum+Strelitzia] or Strelitzia was sister-group to [Phenakospermum+ Ravenala]. Within Strelitzia, S. nicolai is sister to the remaining species.

Cladogram of Strelitziaceae based on DNA sequence data (Prince 2010; Cron & al. 2012)

ZINGIBERACEAE Martinov

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Martinov, Tekhno-Bot. Slovar: 682. 3 Aug 1820 [’Zinziberaceae’], nom. cons.

Amomaceae J. St.-Hil., Expos. Fam. Nat. 1: 154. Jan-Aug 1805, nom. illeg.; Alpiniaceae Link, Enum. Hort. Berol. Alt. 1: 2. 16 Mar-30 Jun 1821; Curcumaceae Dumort., Anal. Fam. Plant.: 56. 1829; Amomales Lindl., Key Bot.: 69. 15-30 Sep 1835

Genera/species c 50/1.270–1.330

Distribution Tropical and southern Africa, Madagascar, islands in the Indian Ocean, South, East and Southeast Asia, Malesia, New Guinea, tropical Australia (especially Queensland), Pacific islands; some species of Renealmia in tropical America (Central America, the West Indies, northern and central South America).

Fossils Fossil fruits, seeds and leaves, Zingiberopsis, of plausible Zingiberaceae are known from Campanian and early Palaeogene layers in North America.

Habit Bisexual, perennial (sometimes giant) herbs, often with a pseudostem consisting of superimposed leaf sheaths. Sometimes with stilt roots. Aromatic. Rhizome and roots usually rich in starch (root cells sometimes with starch nodules).

Vegetative anatomy Phellogen absent. Stem cortex and central tissue with each vascular bundle surrounded by a dense fibre envelope; cortex and central tissue separated by a cylinder of fibres. Secondary lateral growth absent. Vessels present in roots (rarely also in stem). Vessel elements usually with scalariform (sometimes also simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve element nucleus with non-dispersive crystalline protein bodies. Sieve tube plastids P2cs type, with cuneate protein crystals and starch grains, without protein filaments. Nodes? Idioblasts with suberised cell walls and containing abundant aromatic oils and resins. Stegmata-shaped cells with single large silica bodies sometimes present (in Alpinioideae in connection with vascular bundles). Calciumoxalate as rhomboidal, prismatic or tubular crystals (raphides usually absent). Crystal sand present at least in Roscoea. Starch grains usually simple, often flat (sometimes irregular), usually excentric (sometimes compound; rarely spherical or elliptic).

Trichomes Hairs usually absent, when present usually unicellular (on rhizome in Curcuma bicellular; rarely with sunken base; rarely multicellular, furcate to stellate or uniseriate).

Leaves Alternate (distichous), simple, entire, often differentiated into pseudopetiole and pseudolamina, with supervolute ptyxis (laterally rolled-up in bud). Stipules absent; leaf sheath usually open (sometimes closed), usually with a ligule at distal end. Pulvinus usually absent (present in, e.g., Zingiber). Pseudopetiole and mid-vein with cylinder of aerenchyma strands. Vascular bundles abaxial. Venation pinnate-parallelodromous; mid-vein distinct; secondary veins closely spaced, parallel, connected by transverse tertiary veins. Stomata usually tetracytic (sometimes hexacytic or paracytic); guard cells with inner and outer borders non-uniform. Cuticular wax crystalloids usually absent (sometimes present as platelets or rodlets). Epidermis with spherical silica bodies or silica sand. Mesophyll with spherical idioblasts containing ethereal oils and calciumoxalate as druses or prismatic crystals (raphides usually absent). Leaf margin entire.

Inflorescence Flowers on foliar shoots or on special non-photosynthesizing shoots, in terminal usually head- or cone-like (rarely raceme- och spike-like) often branched thyrse consisting of few- to many-flowered (sometimes one-flowered) cincinni subtended by spirally arranged imbricate bracts (bracts rarely absent). Floral prophyll (bracteole) sometimes lateral. Extrafloral nectaries frequent.

Flowers Zygomorphic, often large. Epigyny. Tepals 3+3; outer tepals usually sepaloid, partially connate into a cupulate or asymmetrical spatha-like structure with a lateral longitudinal slit; median inner tepal adaxial, dome-shaped and usually larger than lateral inner tepals; inner tepals petaloid, connate below into a tube (outer and inner tepals in species of Cyphostigma and Elettaria together forming an outgrowth on top of ovary). Gynoecial/septal epigynous nectaries two, inserted at ovary apex; one or three glandular nectaries present at stylar base and staminal bases (sometimes also on staminodia, androecial nectaries). Disc absent.

Androecium Stamen single (median inner adaxial stamen fertile; median outer abaxial stamen absent). Filament of fertile stamen short or long (sometimes absent). Anther dorsifixed, sometimes versatile, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by pores); connective at apex usually prolonged into a pectinate structure of varying shape (in Kaempferia petaloid); placentoid present. Tapetum usually secretory (sometimes amoeboid-periplasmodial), with uninucleate or binucleate cells. Staminodia two or four; two lateral inner staminodia connate into a petaloid labellum; two lateral outer staminodia petaloid or small and reduced (in some genera absent), inserted on both sides of fertile stamen or adnate to labellum; inner tepals adnate to staminodia in Globba and Cautleya; inner tepals and labellum free from each other in Vanoverbergia, Riedelia, Aframomum, Hornstedtia, Zingiber, Amomum, and Cautleya; filament and labellum fused into a tube in Geocharis, Pommereschea, Etlingera, Aulotandra, Stadiochilus, and Globba; lateral staminodia adnate to filament of functioning stamen in Mantisia; staminodia and labellum fused in Siphonochilus.

Pollen grains Microsporogenesis successive. Pollen grains usually inaperturate (sometimes monosulcate or 5–16-pantoporate), shed as monads, bicellular at dispersal. Exine tectate (tectum thin; exine sometimes intectate), with ? infratectum, verrucate, echinate, scabrate, rugulate, striate or smooth, or sometimes absent.

Gynoecium Pistil composed of three connate carpels; median carpel abaxial. Ovary inferior, ab initio trilocular, finally incompletely or entirely trilocular or unilocular. Style single, filiform, hollow, situated in a groove in filament of fertile stamen and between the two anther halves (thecae). Stigma infundibuliform, papillate or non-papillate, Wet type, present above fertile anther. Pistillodium absent.

Ovules Placentation axile (when ovary trilocular); in ‘Globbeae’ parietal with unilocular ovary; in some genera of Zingibereae basal through reduction of septa. Ovules (one to) four to c. 50 per carpel, anatropous or orthotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument (five to) seven to 13 cell layers thick. Inner integument ? cell layers thick. A cylindrical ridge-shaped thickening, ‘micropylar collar’ (‘Mikropylarkragen’), formed from outer integument surrounding micropyle. Hypostase present in many species. Epistase usually present. Nucellar cap sometimes present. Parietal cell formed from archesporial cell. Lateral epidermal cells dividing periclinally. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio helobial; chalazal chamber early degenerating. Endosperm haustoria? Embryogenesis asterad.

Fruit Usually a loculicidal, dry or fleshy capsule (sometimes irregularly dehiscing or indehiscent; in Aframomum a berry; in, e.g., Etlingera a fleshy syncarp).

Seeds Aril usually present. Seed coat testal. Exotesta consisting of fibriform elongate cells and with phytomelan layer on epidermal cell walls. Endotestal cells with silica, usually sclerotic and U-shaped in cross-section (in Globba, Hedychium and their closest relatives thin, not U-shaped). Ethereal oils present in testa. Operculum (formed from inner testal epidermis) present adjacent to radicula (and combined with ‘micropylar collar’). Perisperm well developed, copious, rich in compound starch grains. Chalazosperm absent. Endosperm usually sparse, with aleurone (sometimes with starch grains). Embryo straight (in Elettaria large), without chlorophyll, without lipid bodies, Trillium type. Cotyledon one. Hypocotyl internode short to long. Mesocotyl absent. Cotyledon hyperphyll compact, not assimilating. Coleoptile present or absent. Collar indistinct. Germination cryptocotylar?

Cytology n = 8–14, 17, 21, 25 – Polyploidy and aneuploidy frequently occurring.

DNA Two different indel mutations (insertions), each one comprising one base-pair in plastid 18SrDNA, present in all investigated Zingiberaceae, but not found in other taxa (Soltis & al. 1997). Mitochondrial intron coxII.i3 lost (Kaempferia). Mitochondrial coxI intron present in several genera (including Siphonochilus).

Phytochemistry O-methylated flavonols (kaempferol, myricetin, isorhamnetin, syringetin) flavanones, chalcones, ethereal oils consisting of mono-, di-, tri- and sesquiterpenes, condensed tannins (from proanthocyanidins), proanthocyanidins (prodelphinidin, procyanidin), phenylpropanoids and related curcumins (diarylheptanoids), caffeic acid, quinines, glycosides (e.g. 3-glycoside, 3-rutenoside), aromatic acids and ketones, and shikimic acid- or polyacetate-derived arthroquinones present. Dihydroflavones, alkaloids, steroidal saponins, and cyanogenic compounds rare. Flavone-C-glycosides and ellagic acid not found.

Use Ornamental plants, starch sources (East Indian arrowroot from Curcuma), spices (Aframomum melegueta, Amomum, Boesenbergia, Curcuma, Elettaria cardamomum, Kaempferia, Zingiber officinale, etc.), medicinal plants, dyeing sources (Curcuma), perfumes (Alpinia galanga, A. officinale, Curcuma zedoaria, Hedychium).

Systematics Zingiberaceae are sister to Costaceae. The African-Malagasy Siphonochilus is sister to the remaining Zingiberaceae.

Siphonochiloideae W. J. Kress in W. J. Kress, L. M. Prince et K. J. Williams in Amer. J. Bot. 89: 1691. 2002

1/c 20. Siphonochilus (c 20; tropical and southern Africa, Madagascar). – Rhizome fleshy, vertical. Distichy of leaves perpendicular to rhizome. Inflorescence a raceme. Floral prophylls (bracteoles) absent. Lateral staminodia petaloid, adnate to labellum. Labellum not connate to filament. Anther crest petaloid. Ovary trilocular (sometimes inclompletely so). Placentation axile. Fruit fleshy. n = ?

[Tamijia+[Alpinioideae+Zingiberoideae]]

Tamijioideae W. J. Kress in W. J. Kress, L. M. Prince et K. J. Williams in Amer. J. Bot. 89: 1691. 2002

1/1. Tamijia (1; T. flagellaris; northwestern Borneo). – Rhizome fibrous. Distichy of leaves perpendicular to rhizome. Lateral staminodia petaloid, adnate to labellum. Labellum not connate to filament. Anther crest petaloid. Ovary unilocular. Placentation parietal. Fruit? n = ? – Tamijia is sister to the main clade [Alpinioideae+Zingiberoideae].

[Alpinioideae+Zingiberoideae]

Labellum formed exclusively of the two staminodia in the inner androecial whorl. Placentation sometimes free central. Endosperm without starch.

Alpinioideae (Blume) Hassk., Cat. Hort. Bot. Bogor.: 51. Oct 1844 [‘Alpiniae’]

c 20/790–840. Riedelieae in W. J. Kress, L. M. Prince et K. J. Williams in Amer. J. Bot. 89: 1691. 2002. Siliquamomum (1; S. tonkinense; Vietnam), Siamanthus (1; S. siliquosus; Thailand), Burbidgea (5; Borneo), Pleuranthodium (c 25; New Guinea, tropical Australia), Riedelia (c 60; East Malesia). – Alpinieae (Blume) Orb., Dict. Univ. Hist. Nat. 1: 299. 18 Dec 1841. ’Alpinia’ (200–240; tropical and subtropical Asia and eastwards to islands in the Pacific; non-monophyletic), Aframomum (50–55; tropical Africa, Madagascar, the Mascarene Islands, the Seychelles), Renealmia (c 75; tropical America), ’Amomum’ (c 150; tropical Asia, tropical Australia; non-monophyletic), Elettariopsis (c 10; Southeast Asia, West Malesia), Paramomum (1; P. petaloideum; southwestern China), Plagiostachys (20–25; southern China to West Malesia), ’Etlingera’ (c 100; tropical Asia; non-monophyletic), Hornstedtia (c 50; tropical Asia, tropical Australia), Vanoverberghia (1; V. sepulchrei; the Philippines); unplaced: Aulotandra (5; tropical West Africa, Madagascar), Cyphostigma (1; C. pulchellum; Sri Lanka), Elettaria (7; India, Southeast Asia, West Malesia), Geocharis (6; West Malesia), Geostachys (c 20; Southeast Asia, West Malesia), Leptosolena (2; the Philippines). – Tropical and southern Africa, Madagascar, the Mascarene Islands, the Seychelles, tropical Asia, tropical Australia and islands in the Pacific, tropical America. Rhizome fleshy. Distichy of leaves perpendicular to rhizome. Pseudolamina in Riedelieae with nectar pits (extrafloral nectaries) on abaxial mid-vein. Lateral staminodia in outer whorl very small or absent, never petaloid. Labellum not connate to filament. Anther crest petaloid or absent. Ovary uni- or trilocular. Styloids present in Aframomum. Flexistyly (style changing direction during anthesis) sometimes present in Alpinieae. Placentation axile, parietal or free central. Fruit in Riedelieae silique-like dehiscing with longitudinal slits, in Alpinieae fleshy or indehiscent. n = 11, 12. – Siliquamomum is sister to Riedelieae (Kress & al. 2002).

Zingiberoideae (Blume) Hassk., Cat. Hort. Bot. Bogor.: 49. Oct 1844 [‘Zingibera’]

c 30/480–490. Globbeae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 388, Comm. 290. 17-20 Aug 1842. Globba (c 35; East and tropical Asia), Mantisia (4; tropical Asia). – Gagnepainia clade’ Gagnepainia (3; Southeast Asia), Hemiorchis (3; central Himalayas to Burma). – Caulokaempferiaclade’ Caulokaempferia (12; Himalayas to Southeast Asia). – Zingibereae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 388, Comm. 290. 17-20 Aug 1842. Camptandra (4; West Malesia), Pyrgophyllum (1; P. yunnanense; China), Smithatris (2; Burma, Thailand; in Curcuma?), Hitchenia (3; tropical Asia; in Curcuma?), Stahlianthus (6; northern India, southern and southwestern China, Burma, Thailand; in Curcuma?), Paracautleya (1; P. bhatii; southern India), Curcuma (c 50?; tropical Asia; paraphyletic?),‘Hedychium (c 50?; Madagascar, Himalayas, tropical Asia), Larsenianthus (4; India, Bangladesh, Burma), Nanochilus (2; Sumatra, the Moluccas), Roscoea (17–20; Himalayas, western China), Cautleya (5; Himalayas), Pommereschea (2; southern China, Burma, Thailand), Rhynchanthus (5–6; southwestern China, Burma, West Malesia), Haniffia (2; Peninsular Thailand, the Malay Peninsula), Curcumorpha (1; C. longiflora; eastern Himalayas; in Boesenbergia?), ’Boesenbergia’ (c 80; tropical Asia; non-monophyletic), Kaempferia (c 40; India to southern China and Malesia), Zingiber (c 100; East and Southeast Asia, Malesia to tropical Australia), Cornukaempferia (2; Thailand), Scaphochlamys (c 30; southern Thailand, the Malay Peninsula), Distichochlamys (3; Vietnam); unplaced: Kedhalia (1; K. flaviflora; Kedah Province in the Malay Peninsula), Newmania (2; Vietnam), Haplochorema (3–4; Sumatra, Borneo), Laosanthus (1; L. graminifolius; Laos), Parakaempferia (1; P. synantha; Assam), Stadiochilus (1; S. burmanicus; Burma). – Madagascar, Himalayas, East and tropical Asia, tropical Australia. Rhizome fibrous. Distichy of leaves parallel to rhizome. Lateral staminodia petaloid, adnate to or free from labellum, sometimes connate to filament. Labellum in Zingibereae not connate to filament, in ‘Globbeae’ adnate to filament and forming a tube. Anther crest absent, petaloid, or well developed and wrapped around style (in Zingibereae), or spurred or absent (in ‘Globbeae’). Tapetum sometimes amoeboid-periplasmodial. Pollen grains in Zingiber sulcate. Ovary unilocular or trilocular (sometimes incompletely so). Style with two vascular bundles. Placentation in Zingibereae axile, basal or free columnar, in ‘Globbeae’ parietal. Fruit in Zingibereae fleshy and dehiscent, in ‘Globbeae’ globose and dehiscent. Seeds in Globba and Zingiber carunculate. Exotestal epidermis in ‘Globbeae’ multilayered. Endotesta parenchymatous. Starch grains of perisperm usually compound (in ‘Globbeae’ simple). Embryo in Hedychium long. Steroidal saponins present in Hedychium. n = (8–)10–12(l–14) and higher. – The clade [Globba+Mantisia] forms a sister-group to the remaining Zingiberoideae, and the clade [Gagnepainia+Hemiorchis] is successive sister to [Caulokaempferia+Zingibereae] (Kress & al. 2002).Pommereschea has a parenchymatic endotesta and a long style.

Cladogram (one of 980 most-parsimonious) of Zingiberaceae based on DNA sequence data (Kress & al. 2002).


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