COMMELINALES Dumortier Main Tree, Synapomorphies

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

COMMELINALES Mirb. ex Bercht. et J. Presl

Habit Usually bisexual (sometimes monoecious or andromonoecious, rarely polygamomonoecious), usually perennial (rarely annual) herbs. Usually somewhat succulent, sometimes twining, very rarely epiphytic. Bulb, corm or tuber rarely present. Often aquatic or helophytic.

Vegetative anatomy Mycorrhiza not found. Phellogen absent. Primary vascular tissue cylinder of bundles, or two or several cylinders, or scattered bundles. Secondary lateral growth absent. Vessels in roots, and sometimes stems and/or leaves. Vessel elements usually with scalariform (sometimes simple or reticulate) perforation plates; lateral pits scalariform or alternate, simple or bordered pits. Imperforate tracheary xylem elements tracheids with simple pits. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals, without starch or protein filaments. Nodes usually multilacunar with several leaf traces. Mucilage ducts and chambers with cells containing calciumoxalate raphides. Latex rarely present. Tanniniferous (sometimes lignified) idioblasts sometimes present. Silica bodies usually absent. Prismatic calciumoxalate crystals rarely present.

Trichomes Hairs unicellular or multicellular, usually uniseriate (rarely multiseriate), simple or branched; tricellular glandular microhairs present in most clades.

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, usually linear, sometimes equitant (ensiform), rarely subulate, with conduplicate, involute, convolute or supervolute (rarely plicate) ptyxis, sometimes differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath closed, open or absent (rarely reduced to ligule). Colleters sometimes present. Venation (pinnate-)parallelodromous, sometimes acrodromous. Stomata paracytic, tetracytic or hexacytic. Cuticular wax crystalloids as aggregated rodlets (Strelitzia type), non-aggregated rodlets or absent. Epidermis thick, sometimes with idioblasts containing silica bodies. Mesophyll often with mucilaginous idioblasts or mucilage canals containing calciumoxalate raphides (mesophyll rarely with tanniniferous idioblasts). Leaf margin entire.

Inflorescence Terminal or axillary, panicle, thyrse, or corymb, often composed of cincinni, sometimes raceme-, head- or spike-like (flowers rarely solitary). Bracts sometimes large, sometimes boat-shaped. Floral prophylls (bracteoles) sometimes lateral or absent.

Flowers Actinomorphic or zygomorphic. Hypogyny, epigyny or half epigyny. Tepals usually 3+3 (sometimes 2+2, rarely 3+2); outer tepals usually with imbricate (rarely valvate) aestivation, sepaloid or petaloid, free or more or less connate; inner tepals with usually imbricate (rarely valvate) aestivation, usually petaloid (rarely sepaloid), thin, often stipitate, free or more or less connate (sometimes into tube); when zygomorphic, then median inner tepal often abaxial. Nectary absent, or septal nectaries inserted below insertion point of tepals or present on staminodia. Disc absent.

Androecium Stamens 3+3, two to four of which often staminodial, or three fertile stamens and no staminodium (sometimes one fertile stamen and two staminodia, or one stamen and no staminodium). Filaments glabrous or entirely or partially covered with long and usually moniliform hairs, free from each other and from tepals. Anthers basifixed to dorsifixed, sometimes versatile, tetrasporangiate, introrse, latrorse, or extrorse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum usually amoeboid-periplasmodial (sometimes secretory). Staminodia absent or alternating with stamens (antesepalous or antepetalous), or inserted on only one (anterior or posterior) side of flower, and stamens on opposite side; female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (sometimes disulcate to tetrasulcate, or monosulculate to trisulculate, rarely inaperturate, diporate, triporate, heptaporate or octoporate), usually shed as monads (rarely tetrads or polyads), bicellular or tricellular at dispersal. Exine tectate or intectate, usually with columellate (sometimes acolumellate) infratectum, verrucate, rugulate, spinulate or granulate.

Gynoecium Pistil composed of (one to) three connate carpels. Ovary superior, inferior or semi-inferior, unilocular or entirely or partially bilocular or trilocular. Style single, simple, straight or curved, usually narrow. Stigma simple, small or large, punctate, capitate, penicillate, triangular or somewhat trilobate, papillate or non-papillate, Dry or Wet type. Male flowers often with pistillodium.

Ovules Placentation usually axile (rarely parietal or basal). Ovules one to more than 50 (to more than 100) per carpel, orthotropous, hemianatropous or campylotropous (sometimes hypotropous, pleurotropous, anatropous or irregular), ascending or pendulous, apotropous, epitropous, bitegmic, crassinucellar or weakly crassinucellar. Micropyle endostomal or bistomal (sometimes exostomal). Funicular obturator sometimes present. Parietal cell usually formed from archesporial cell (sometimes further dividing forming parietal tissue). Nucellar cap sometimes present. Periclinal divisions sometimes taking place in megasporangial epidermis (then parietal cell not formed). Megagametophyte monosporous, usually Polygonum type (rarely disporous, Allium type, or tetrasporous, Adoxa, Oenothera or Scilla types). Synergids sometimes with a filiform apparatus. Antipodal cells ephemeral, not proliferating. Endosperm development nuclear or helobial. Cell wall formation in small chalazal chamber preceding that in large micropylar chamber. Endosperm haustoria micropylar or absent. Embryogenesis asterad or onagrad.

Fruit A loculicidal or poricidal capsule (rarely irregularly dehiscent or indehiscent, a nut, or a schizocarp with nut-like mericarps), a berry-like capsule or berry.

Seeds Aril sometimes present. Seed coat testal-tegmic. Exotesta sometimes with pigmented outer epidermis, sometimes collapsed, sometimes with thick cell walls. Endotestal cells often with silica bodies. Tegmic cells sometimes compressed, sometimes degenerating or crushed. Perisperm not developed. Endosperm copious, starchy, often also with lipids and aleurone. Embryo small, straight, situated opposite hilum (often near micropyle), usually well differentiated, often covered with discoid or conical embryostega, testal operculum, usually surrounded by micropylar collar, without chlorophyll, Xyris-Scirpus or Trillium type. Cotyledon usually one (sometimes two, one of which rudimentary). Cotyledon hyperphyll elongate or compact, often assimilating. Hypocotyl internode short to long, or absent. Mesocotyl usually absent. Coleoptile often present. Collar rhizoids usually present. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, syringetin), O-methylated flavonols, 6-hydroxyflavonoids (6-hydroxyluteolin), flavone-C-glycosides, cyanidin, cyanidin-3,7,3’-triglycoside, delphinidin, proanthocyanidins (prodelphinidins), phenolic acids, chelidonic acid, tyrosine-derived cyanogenic compounds, phenylphenalenones (perinaphthenones), arylphenalenones, polyamines, and fructans present. Ferulic or diferulic acids and p-coumaric acids often present in cell walls. Alkaloids and steroidal saponins rare. Ellagic acid not found.

Stevens (2001 onwards) lists the following potential synapomorphies of the clade [Philydraceae+[Haemodoraceae+Pontederiaceae]]: absence of silica bodies; presence of styloids; perianth with tanniniferous cells; perianth persistent in fruit; and placentae containing sclereids. Haemodoraceae and Pontederiaceae have the following characters in common (Stevens 2001 onwards): ectexine not tectate or columellate; endothecial cells with basal plates; and presence of phenylphenalenones (also found in some clades of Cannales).

Ephemeraceae Batsch, Tab. Affin. Regni Veg.: 125. 2 Mai 1802 [‘Ephemera’], nom. illeg.; Tradescantiaceae Salisb. in Trans. Linn. Soc. London 8: 9. 1807 [‘Tradescanteae’]; Ephemerales Burnett, Outl. Bot.: 1153. Jun 1835 [‘Ephemerinae’], nom. illeg.; Cartonemataceae Pichon in Notul. Syst. (Paris) 12: 219. Feb 1946, nom. cons.

Distribution Tropical and subtropical regions, with the largest diversity in Africa, South Asia, Mexico and northern Central America; some species in temperate East Asia, eastern and southern North America and Australia.

Fossils Leaves and fruits have been found in mid-Miocene layers in Kenya and interpreted as Pollia.

Habit Usually bisexual (often monoecious or andromonoecious, rarely polygamomonoecious), usually perennial (rarely annual) herbs. Usually somewhat succulent, sometimes twining, very rarely epiphytic. Bulb rarely present. Nodes swollen.

Vegetative anatomy Mycorrhiza absent. Roots fibrous or tuberous. Phellogen absent. Cortex usually without vascular tissue (cortical vascular bundles usually absent; present in Cartonema). Primary vascular tissue as scattered bundles. Central vascular cylinder surrounded by endodermis-like envelope; longitudinal vascular bundles arranged in a typical pattern and connected only at nodes. Secondary lateral growth absent. Vessels in root, stem and leaves. Vessel elements with scalariform or reticulate (Cartonematoideae), or simple (Commelinoideae) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Raphide canals (mucilage canals) consisting of cells with calciumoxalate raphides (usually absent in Cartonematoideae). Silica bodies usually absent (present in some species).

Trichomes Hairs multicellular, usually uniseriate (in Palisota multiseriate, with some cells thin-walled and sometimes branched); tricellular glandular microhairs present in most genera (absent in Cartonematoideae).

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, often linear, with usually convolute (supervolute; sometimes involute, rarely plicate) ptyxis. Stipules absent; leaf sheath closed; ligule absent. Venation parallelodromous; midvein distinct; few primary veins connected to thin secondary and tertiary veins. Stomata tetracytic, hexacytic or paracytic. Cuticular wax crystalloids as longitudinally aggregated rodlets (Strelitzia type). Epidermis thick, sometimes with silica bodies. Mesophyll with mucilaginous idioblasts or mucilaginous canals containing calciumoxalate raphides (absent in Cartonematoideae). Leaf margin entire.

Inflorescence Terminal and/or axillary, principally cymose paniculate thyrse composed of helicoid partial inflorescences (cincinni), in Cartonematoideae raceme- or spike-like (flowers sometimes solitary), in some species surrounded by large bracts, spathae. Bracts sometimes boat-shaped. Floral prophyll (bracteole) usually median (prophylls in Dichorisandra lateral).

Flowers Actinomorphic or zygomorphic (transversally so i Dichorisandra; asymmetric in, e.g., Cochliostema). Hypogyny. Tepals usually 3+3 (rarely 3+2); outer tepals with imbricate aestivation, usually sepaloid or scale-like (sometimes petaloid), free or connate below; inner tepals with imbricate aestivation, petaloid, thin, often stipitate, usually early caducous, free or connate at base (in some species connate into a tube); when zygomorphic, then usually median inner tepal abaxial. Nectaries usually absent (possibly present on staminodia in species of Commelina). Septal nectaries absent. Disc absent. Enantiostyly present in Murdannia.

Androecium Stamens usually 3+3 (rarely fewer), two or three (rarely four) of which often staminodial (rarely three fertile stamens and no staminodia); some stamens often strongly yellow, showy, attracting and with fewer pollen grains than in remaining stamens. Filaments glabrous or often entirely or partially covered with long and usually moniliform hairs, free from each other and from tepals. Anthers basifixed or dorsifixed, sometimes versatile, tetrasporangiate, introrse, latrorse, or extrorse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores); connective sometimes slightly prolonged. Tapetum amoeboid-periplasmodial; tapetal cell contents early invading anther cavity. Tapetal cells often with calciumoxalate raphides. Endothecium present. Staminodia absent or alternating with stamens (antesepalous or antepetalous), or inserted on one (anterior or posterior) side of flower, and stamens on opposite side; female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (rarely inaperturate or di-, tri- or tetrasulcate), shed as monads, bicellular or tricellular at dispersal. Exine tectate, with columellate infratectum, verrucate, spinulate or granulate.

Gynoecium Pistil composed of usually three connate carpels (sometimes two by degeneration of dorsal carpel); median carpel abaxial. Ovary superior, usually bilocular or trilocular (rarely unilocular; sometimes unilocular at apex and trilocular in lower part). Style single, simple, straight or curved, usually narrow. Stigma simple, small or large, punctate, capitate, penicillate, triangular or somewhat trilobate, papillate or non-papillate, Dry or Wet type. Male flowers often with pistillodium.

Ovules Placentation usually axile (rarely parietal or basal). Ovules one to c. 20 (to more than 50; in Cartonematoideae two) per carpel, orthotropous, hemianatropous or campylotropous (rarely anatropous), ascending, bitegmic, weakly crassinucellar. Micropyle usually endostomal (sometimes exostomal). Outer integument three to seven (to ten) cell layers thick. Inner integument two cell layers thick. Nucellar cap one or two cell layers thick. Parietal cell usually formed from archesporial cell. Parietal tissue one or two cell layers thick. Periclinal divisions sometimes taking place in megasporangial epidermis (parietal cell not formed). Megagametophyte monosporous, usually Polygonum type (rarely disporous, Allium type, or tetrasporous, Adoxa, Oenothera or Scilla types). Antipodal cells usually early degenerating (in Tinantia persistent). Endosperm development ab initio nuclear. Endosperm haustorium chalazal. Embryogenesis asterad.

Seeds Aril present in some genera. Operculum formed by testa (and combined with micropylar collar). Seed coat usually endotestal-exotegmic. Exotesta thin. Endotestal cells usually with silica bodies. Endotegmen thin. Perisperm not developed. Endosperm copious, starchy; starch grains usually complex. Suspensor absent. Embryo small, straight, situated opposite hilum, usually well differentiated (in Cartonematoideae little differentiated), covered by discoid or conical operculum, embryotega, usually surrounded by a micropylar collar (absent in Cartonema), without chlorophyll (non-photosynthesizing), Xyris-Scirpus type. Cotyledon usually one (sometimes two, one of which rudimentary). Cotyledon hyperphyll elongate (directed downwards) or compact, not assimilating. Hypocotyl internode long, short or absent. Mesocotyl usually absent (present in Cartonema). Coleoptile often present. Collar rhizoids usually present (absent in Cartonema). Germination phanerocotylar.

Cytology n = 4, 6, 7, 9–17, 20, 29 (Commelinoideae); n = 12 (Cartonematoideae) – Polyploidy and aneuploidy occurring.

Phytochemistry Flavonols, 6-hydroxyluteolin, flavone-C-glycosides, cyanidin-3,7,3’-triglycoside, and phenolic acids present. Alkaloids, steroidal saponins, and tyrosine-derived cyanogenic compounds rare. Ellagic acid and chelidonic acid not found.

Systematics Commelinaceae are probably sister-group to Hanguanaceae. Cartonema (Australia) and Triceratella (Africa) together form a sister to the remaining Commelinaceae.

Cartonematoideae (Pichon) Faden ex G. C. Tucker in J. Arnold Arbor. 70: 99. 10 Jan 1989

2/12. Cartonema (11; tropical Australia), Triceratella (1; T. drummondii; Zimbabwe). – Cortical vascular bundles present. Stem collenchyma absent. Vessel elements with scalariform or reticulate perforation plates. Silica bodies absent. Raphide canals present adjacent to veins or absent. Stomata paracytic or tetracytic. Microglands absent; normal glandular hairs frequent. Inflorescence racemose or with a single flower per cincinnus. Flowers sessile, yellow. Ovules two per carpel. Fruit a berry. Main part of testa caducous. Operculum without collar. Collar short. Collar rhizoids absent. Mesocotyl present. Radicula stout. n = 12.

Commelinoideae Eaton, Bot. Dict., ed. 4: 27. Apr-Mai 1836 [‘Commelineae’]

35/645–655. Palisota (18); Commelineae Dumort., Anal. Fam. Plant.: 55. 1829. Murdannia (50–55; tropical and subtropical regions on both hemispheres), Anthericopsis (1; A. sepalosa; tropical East Africa), Buforrestia (3; B. candolleana: French Guiana, Suriname; B. mannii, B. obovata: tropical West and Central Africa), Floscopa (c 20; tropical and subtropical regions on both hemispheres), Stanfieldiella (4; S. axillaris, S. brachycarpa, S. imperforata, S. oligantha; tropical Africa), Commelina (c 170; tropical and subtropical regions on both hemispheres), Polyspatha (3; P. hirsuta, P. oligospatha, P. paniculata; tropical West Africa), Aneilema (68; tropical and southern Africa, Madagascar, Yemen, few species in tropical Asia to tropical Australia and islands in southwestern Pacific; one species, A. brasiliense; in tropical South America), Pollia (17; tropical regions in the Old World, tropical Australia; one species, P. americana, in Panamá). – Tradescantieae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 406. 17-20 Aug 1842. Spatholirion (6; S. calcicola, S. decumbens, S. elegans, S. longifolium, S. ornatum, S. puluongense; southern China, Southeast Asia), Cochliostema (2; C. odoratissimum, C. velutinum; Nicaragua to Ecuador), Geogenanthus (3; G. ciliatus, G. poeppigii, G. rhizanthus; tropical South America); Tinantia (13–14; Texas to tropical South America), Thyrsanthemum (3; T. floribundum, T. goldianum, T. macrophyllum; Mexico), Weldenia (1; W. candida; Mexico, Guatemala), Elasis (2; E. hirsuta: Ecuador; E. guatemalensis: Central America), Callisia (c 20; southern United States to Argentina), Tradescantia (c 80; North to South America), Dichorisandra (c 30; Central America, the West Indies, tropical South America), Siderasis (1; S. fuscata; southeastern Brazil), Amischotolype (15; tropical regions in the Old World), Coleotrype (9; tropical and southern Africa, Madagascar), Belosynapsis (6; B. ciliata, B. epiphytica, B. kawakamii, B. kewensis, B. moluccana, B. vivipara; Madagascar, tropical Asia from India to New Guinea), Cyanotis (c 50; tropical and subtropical regions of the Old World, from Africa to northern Australia). – Unplaced Commelinoideae Aetheolirion (1; A. stenolobium; Thailand), Dictyospermum (5; D. conspicuum, D. humile, D. montanum, D. ovalifolium, D. ovatum; India, Sri Lanka to New Guinea), Gibasoides (1; G. laxiflora; Mexico), Matudanthus (1; M. nanus; Mexico), Plowmanianthus (5; P. dressleri, P. grandifolius, P. panamensis, P. perforans, P. peruvianus; Panama, Amazonas), Pseudoparis (3; P. cauliflora, P. monandra, P. tenera; Madagascar), Sauvallea (1; S. blainii; Cuba), Streptolirion (1; S. volubile; eastern Himalayas to Southeast Asia and the Korean Peninsula), Tapheocarpa (1; T. calandrinioides; northeastern Queensland), Tricarpelema (8; tropical Asia; one species, T. africanum, in Cameroon, Gabon and Equatorial Guinea). – Tropical, subtropical, temperate parts of North America and East Asia. Cortical vascular bundles absent. Stem collenchyma present. Vessels present also in stem and leaves. Vessel elements with simple perforation plates. Stem with narrow cortex and endodermis-like envelope surrounding vascular bundles connected to one another only at nodes. Silica bodies sometimes present. Raphide canals present between veins. Stomata tetracytic, hexacytic, and other types. Tricellular microglands present. Petals sometimes connate into a tube. Stamens one to six. Filaments often densely beset with uniseriate hairs. Anthers often procidal. Staminodia often two to four, attracting pollinators. Pollen grains usually with calciumoxalate raphides. Polar nuclei early fusing. Seed operculum (embryotega) testal, with one micropylar collar. Collar roots present. Mesocotyl absent. Cyanidin-3,7,3’-triglucoside present. – Palisota is sister to the remaining Commelinoideae in the study by Evans & al. (2003).

Cladogram of Commelinaceae based on DNA sequence data (Evans & al. 2003).

Xiphidiaceae Dumort., Anal. Fam. Plant.: 59, 61. 1829 [‘Xiphideae’, ‘Xiphidieae’, ‘Xiphineae’]; Haemodorales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 9. Sep-Oct 1835 [‘Haemodoraceae’]; Wachendorfiaceae Herb., Amaryllidaceae: 48. late Apr 1837; Dilatridaceae M. Roem., Handb. Allg. Bot. 3: 476. 1840 [’Dilatrideae’]; Conostylidaceae (Benth.) Takht., Sist. Magnoliof. [Systema Magnoliophytorum]: 313. 24 Jun 1987

Distribution South Africa, New Guinea, Australia, Tasmania, eastern and southeastern North America, Central America, northern South America, with their largest diversity in Australia.

Habit Bisexual, perennial herbs. Tuberous stem (Pyrrorhiza, Tribonanthes) or bulb (Haemodorum) and often stolons. Roots and subterranean stems often intensely red to reddish-brown.

Vegetative anatomy Mycorrhiza absent. Phellogen absent. Primary vascular tissue a single cylinder of bundles, or two or several bundle cylinders, or scattered bundles. Stem vascular bundles numerous, collateral to amphivasal, surrounded by a sclerified cylinder. Secondary lateral growth absent. Vessels at least in root metaxylem (in Lachnanthes also in stem). Vessel elements with simple or scalariform perforation plates; lateral pits scalariform or alternate, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids with simple pits. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Coloured latex present in some representatives. Tanniniferous lignified idioblasts present in Conostylidoideae. Calciumoxalate raphides usually numerous. Styloids? Silica bodies usually absent (often present in Conostylis).

Trichomes Hairs unicellular or multicellular, narrowing, pilate, uniseriate or branched, usually with multicellular base.

Leaves Alternate (distichous), simple, entire, linear, often ensiform, isobifacial, sometimes subulate (in Tribonanthes tubular), often coriaceous, with conduplicate or plicate ptyxis. Stipules absent; leaf sheath short. Venation parallelodromous. Stomata paracytic. Cuticular wax crystalloids small or non-orientated. Mesophyll in Dilatris with mucilaginous idioblasts usually containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, usually panicle, thyrsoid, racemoid, head-like or corymb, usually compound, with helicoid partial inflorescences (flowers in Tribonanthes uniflora solitary). Extrafloral nectaries sometimes present.

Flowers Actinomorphic or zygomorphic (sometimes transversally zygomorphic, sometimes split-monosymmetric; in Anigozanthus somewhat oblique), sometimes large. Epigyny, half epigyny or hypogyny (probably secondary). Tepals 3+3, membranous to thick, externally usually with numerous narrowing, pilate or dendritic hairs (sometimes glabrous), usually with imbricate aestivation (posterior outer tepal median), free or connate only at base (tepals in Anigozanthos, Blancoa, Conostylis six in a single whorl, usually with valvate aestivation, connate and tubular); tanniniferous cells? Septal nectaries usually present below insertion point of tepals (absent in Phlebocarya and Xiphidium). Disc absent. Mucilaginous chamber present in flowers of Dilatris. Enantiostyly present in Wachendorfia.

Androecium Stamens three antepetalous (outer staminal whorl absent in most Haemodoroideae; in Pyrrorhiza two staminodia and one fertile stamen; in Schiekia two staminodia and four fertile stamens) or 3+3 (Conostylidoideae). Filaments subulate or flattened, free, often adnate to inner tepals or perianth tube, respectively. Anthers basifixed, subbasifixed or dorsifixed, sometimes versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective sometimes with appendage. Tapetum usually amoeboid-periplasmodial (sometimes secretory). Tapetal cells often with calciumoxalate raphides. Staminodia in Pyrrorhiza and Schiekia two.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate (Haemodoroideae) or di- or triporate or hepta- or octoporate (Conostylidoideae), heteropolar, shed as monads, bicellular at dispersal. Pollen grains sometimes without starch. Exine usually two-layered (sometimes one- or three-layered), intectate, without foot layer, verrucate (Haemodoroideae, rarely foveolate) or rugulate (Conostylidoideae).

Gynoecium Pistil composed of usually three connate carpels (in Barberetta only one fertile). Ovary inferior, semi-inferior or superior (probably secondarily), usually trilocular (in Phlebocarya trilocular at base and unilocular at apex; in Barberetta unilocular due to reduction of two carpels). Style single, subulate or flattened, simple or trifid, with stylar canal. Stigma single, round, or stigmas three, papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation usually axile (in Phlebocarya basal); placentae swollen. Ovules one to more than 50 per carpel, anatropous (hemianatropous?) or orthotropous, hypotropous or pleurotropous, or irregular, bitegmic, crassinucellar. Micropyle usually endostomal (sometimes exostomal). Outer integument usually two cell layers thick. Inner integument usually two cell layers thick. Parietal cell formed from archesporial cell. Nucellar cap sometimes present. Megagametophyte monosporous, Polygonum type. Endosperm development helobial. Endosperm haustoria? Embryogenesis probably asterad (or onagrad).

Fruit A loculicidal to apically poricidal capsule (in some species indehiscent; in Phlebocarya a nut; in one species of Anigozanthos a schizocarp with three nut-like mericarps).

Seeds Aril absent. Caruncle present in Wachendorfia. Testal cells elongate, more or less thin-walled. Outer exotestal epidermis sometimes with brown or black pigments. Exotesta collapsed in Conostyloideae. Tegmic cells compressed, brown, sometimes elongate. Operculum absent. Perisperm not developed. Endosperm copious, with outer layer of thick-walled pitted cells containing lipids and aleurone, and inner layer of thin-walled unpitted cells containing starch; starch grains usually simple (sometimes compound). Embryo small, present near micropyle, Trillium type, chlorophyll? Cotyledon one. Hypocotyl internode short to long. Mesocotyl? Cotyledon hyperphyll elongate, assimilating, or compact, not assimilating. Coleoptile absent (sometimes with a median lobe of cotyledon sheath). Collar rhizoids? Germination?

Cytology n = 4–8, 11, 14, 16, 21, 28 (Conostylidoideae); n = 12, 15, 19–21, 24 (Haemodoroideae)

Phytochemistry O-methylated flavonols (kaempferol, quercetin), proanthocyanidins (cyanidin, prodelphinidin), chelidonic acid, phenylphenalenones (perinaphthenones), arylphenalenones, and fructans present. Diferulic acids and p-coumaric acids in cell walls. Ellagic acid, saponins, and cyanogenic compounds not found. Alkaloids?

Systematics Haemodoraceae may be sister-group to Pontederiaceae. There are two distinct main clades in Haemodoraceae (Hopper & al. 2009): Haemodoroideae and Conostylidoideae, the latter endemic in Southwest Australia.

Haemodoroideae Arn., Botany: 133. 9 Mar 1832 [’Haemodoreae’]

8/34. Dilatris (4; D. corymbosa, D. ixioides, D. pillansii, D. viscosa; Western Cape), Haemodorum (20; southwestern, northern and eastern Australia, Tasmania), Lachnanthes (1; L. caroliniana; Nova Scotia and Massachusetts to Louisiana and Florida, Cuba), Barberetta (1; B. aurea; Eastern Cape, KwaZulu-Natal), Wachendorfia (4; W. brachyandra, W. multiflora, W. paniculata, W. thyrsiflora; Western and Eastern Cape), Schiekia (1; S. orinocensis; northern South America, Brazil, Bolivia), Xiphidium (2; X. caeruleum: southern Mexico Central America, the West Indies, tropical South America; X. xanthorrhizon: western Cuba, Isla de la Juventud), Pyrrorhiza (1; P. neblinae; Venezuela). – Tropical, subtropical and warmtemperate regions, southern Africa, in Southeast Asia only West of Wallace’s line, absent from southern South America. Bulb sometimes present. Roots red. Hairs with distinct basal cells. Zygomorphic flowers resupinate (median inner tepal abaxial, median carpel adaxial). Hypogyny (secondary). Perianth sometimes differentiated. Adaxial outer tepal in Wachendorfia and Schiekia adnate at base to the lateral/abaxial outer tepals and the two lateral/adaxial inner tepals. Flowers often enantiostylous. Supralocular septal nectaries present in Dilatris. Stamens usually three (antepetalous; outer staminal whorl usually absent; in Pyrrorhiza a single stamen; staminodia sometimes present). Pollen grains monosulcate. Seeds often flattened, finely hairy or winged. Cotyledon not photosynthesizing. Hypocotyl short or absent. n = 12, 15, 19–21, 24. Tannins absent.

Conostylidoideae Lindl., Veg. Kingd.: 153. Jan-Mai 1846 [‘Conostyleae’]

6/62–70. Tribonanthes (6–7; T. australis, T. brachypetala, T. longipetala, T. minor, T. purpurea, T. violacea; southwestern Western Australia), Phlebocarya (3; P. ciliata, P. filifolia, P. pilosissima; southwestern Western Australia), Blancoa (1; B. canescens; southwestern Western Australia), Conostylis (40–45; southwestern Western Australia), Anigozanthos (11–12; southwestern Western Australia), Macropidia (1; M. fuliginosa; western Western Australia). – Mainly southwestern Australia. Tanniniferous lignified idioblasts present. Silica sand present. Epidermal cell walls sometimes thickened. Hairs branched. Leaf margin sometimes spiny. Inner tepals often connate (in Anigozanthus six tepals connate into a tube). Flowers usually hairy, sometimes pseudomonocyclic, valvate, zygomorphic. Supralocular septal nectaries usually present. Stamens 3+3. Filaments sometimes adnate to inner tepals. Pollen grains 2–3-porate or 7–8-porate. Placentae with tanniniferous cells. Fruit sometimes a schizocarp or indehiscent. Seeds sometimes ridged. Cotyledon photosynthesizing. Hypocotyl present. Radicula well developed. n = 4–8, 11, 14, 16, 21, 28.

Cladogram of Haemodoraceae based on DNA sequence data (Hopper & al. 2009).

Vegetative anatomy Phellogen absent. Stem cortex thin, with scattered vascular bundles. Endodermis distinct. Central tissue of stem starchy and with fibrous vascular bundles. Primary vascular tissue as scattered closed bundles. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements probably tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Tanniniferous idioblasts and mucilage canals present. Silica bodies present. Raphides rare.

Trichomes Hairs multicellular, branched, with sunken uniseriate base (frequent especially on younger individuals).

Leaves Alternate (spiral), simple, entire, sometimes linear, with supervolute ptyxis, differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath open. Venation pinnate-parallelodromous, tessellate, with distinct midvein and numerous transversal secondary veins. Stomata tetracytic. Cuticular waxes absent. Silica bodies present in endodermis, abaxial hypodermis and mesophyll in some species. Epidermis without silica bodies. Mesophyll with tanniniferous idioblasts. Leaf margin entire.

Inflorescence Terminal, panicle with spike-like partial inflorescences consisting of few-flowered monochasia. Floral prophyll (bracteole) absent.

Flowers Actinomorphic, small. Pedicel absent. Hypogyny. Tepals 3+3, pseudomonocyclic, sepaloid, bract-like, persistent, connate at base. Septal (gynoecial) nectary tripartite, present in male flowers; basal-adaxial staminodial nectaries present in female flowers. Disc absent (male floral receptacle terminating in an accumulation of fleshy structures).

Androecium Stamens 3+3. Filaments filiform, wider at base, almost free (connate at base), adnate to tepals at base. Anthers basifixed to subbasifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial. Female flowers with 3+3 reduced staminodia, three inner ones nectar-secreting (nectaries).

Pollen grains Microsporogenesis successive. Pollen grains inaperturate, shed as monads, ?-cellular at dispersal. Pollen grains without starch. Exine tectate, with columellate? infratectum, spinulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, unilocular above and trilocular below, with mucilaginous hairs on inner side. Style single, simple, very short and wide, or absent. Stigma deeply trilobate, type? Male flowers with reduced pistillodium consisting of three stigmatic lobes and three basal nectaries.

Ovules Placentation basal-axile. Ovule one per carpel, orthotropous to hemianatropous, pendulous, bitegmic, tenuinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator funicular. Epidermal cells anticlinally elongate. Parietal cell not formed (parietal tissue absent). Suprachalazal zone massive. Megagametophyte? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Seed enclosing placenta during development. Seed coat testal(-tegmic). Testa approx. five cell layers thick. Mesotesta sclerified. Endotesta sclerified, with thickened inner periclinal cell walls. Tegmen collapsing and degenerating, with two layers of crossing fibres. Perisperm not developed. Endosperm copious, starchy, surrounded by a layer of aleurone-rich cells. Embryo small, well differentiated, without chlorophyll. Cotyledon one, compact, not photosynthesizing. Hypocotyl internode? Cotyledon hyperphyll? Coleoptile? Collar rhizoids? Radicula well developed. Germination? First leaf of seedling a cataphyll.

Phytochemistry Insufficiently known. Cyanidin and proanthocyanidins (prodelphinidins) present. Ferulic acid present in cell walls. Flavonols not found.

Link, Enum. Hort. Berol. Alt. 1: 5. 16 Mar-30 Jun 1821 [’Philhydrinae’], nom. cons.

Distribution India, the Andaman Islands, East and Southeast Asia to New Guinea, southwestern, northern and eastern Australia, Guam, with their highest diversity in Australia.

Habit Bisexual, perennial herbs. Philydrella has a starch-storing corm. Roots fibrous.

Vegetative anatomy Phellogen absent. Stem cortex and stem centre in Philydrum with vascular bundles and in between with lignified sclerenchymatous tissue containing small collateral or semiconcentric bundles. Secondary lateral growth absent. Vessels present in root (and sometimes stem) metaxylem. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids, simple pits? Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Tanniniferous cells frequent. Laticifers absent. Idioblasts with calciumoxalate styloids abundant. Silica bodies absent.

Trichomes Hairs multicellular, uniseriate, with long terminal cell, densely spaced and woolly; glandular hairs small, sunken.

Leaves Alternate (usually spiral, lower leaves usually distichous), usually isobifacial-equitant (ensiform) with margin facing stem (not in Philydrella), simple, entire, usually linear (in Philydrella subulate), with conduplicate ptyxis. Stipules and ligule absent; leaf sheath short, open. Venation parallelodromous, with numerous primary veins connected to transversal secondary veins. Stomata paracytic (sometimes almost tetracytic). Cuticular wax crystalloids? Mesophyll with cells containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, simple or compound spike in axil of spathe-like bract, sometimes partially adnate to flowers.

Flowers Zygomorphic. Hypogyny or almost hypogyny. Tepals 2(–3)+2(–3), petaloid, usually partially connate (rarely free; all tepals in Helmholtzia connate in lower part); median outer tepal larger than two lateral inner tepals (in Philydrum two inner tepals very small); two lateral outer tepals and median adaxial inner tepal connate into an upper lip; median abaxial outer tepal forming a lower lip; two lateral inner tepals small and usually free (in Philydrella adnate to stamen); tanniniferous cells? Nectaries present in some representatives. Septal nectaries absent. Disc absent. Enantiostyly present.

Androecium Stamen single (probably corresponding to median abaxial inner stamen). Filament wide, flattened, usually free from (sometimes adnate to) median abaxial inner tepal. Anther basifixed to dorsifixed, versatile, peltate, tetrasporangiate, usually introrse to latrorse (in Philydrella curved and hence extrorse; in Philydrum spirally twisted), longicidal (dehiscing by longitudinal slits). Placentoid present. Tapetum secretory, with uninucleate to quadrinucleate cells. Tapetal cells often with calciumoxalate raphides. Staminodia usually absent (in Philydrum rarely a small lateral staminodium from outer staminal whorl).

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, trisulcate, monosulculate, trisulculate or zonoaperturate, shed as monads, tetrads (Philydrum) or polyads, bicellular at dispersal, with calciumoxalate raphides. Exine tectate or semitectate, with columellate infratectum, reticulate or microreticulate.

Gynoecium Pistil composed of three connate carpels; median carpel abaxial, often smaller than lateral carpels. Ovary superior or almost so, trilocular, or unilocular in upper part and trilocular in lower part (Philydrum). Style single, simple. Stigma large, usually capitate (sometimes trilobate), papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (when ovary trilocular; in Philydrum intrusively parietal and ovary unilocular in upper part). Ovules c. 15 to more than 100 per ovary (Philydrum) or c. 15 to c. 50 per carpel (Philydrella, Helmholtzia), anatropous, apotropous or epitropous, pleurotropous, bitegmic, weakly crassinucellar. Micropyle ?-stomal. Outer integument one or two cell layers thick. Inner integument two or three cell layers thick. Obturator funicular. Hypostase present. Parietal cell formed from archesporial cell; parietal tissue sometimes two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development helobial of an aberrant type: small chalazal endosperm chamber with rapid cell division as compared to large micropylar chamber, which develops into nutrient tissue. Endosperm haustorium chalazal. Embryogenesis variation of onagrad.

Fruit Usually a loculicidal capsule (rarely irregularly dehiscent; fruit in Helmholtzia berry-like, leathery).

Seeds Aril absent. Seed sometimes with chalazal hood-shaped structure. Exotestal outer epidermis at micropylar end forming caruncle-like prolongation formed from outer integument. Exotestal cells thick-walled. Endotesta? Exotegmen? Endotegmen tanniniferous. Operculum formed from tegmen (and combined with micropylar collar?). Perisperm not developed. Endosperm copious, proteinaceous, oily and starchy, and with crystalline aleurone bodies; starch grains large elliptic to bean-shaped, and small isodiametric, respectively. Suspensor almost undeveloped. Embryo straight, well differentiated, Trillium type, with chlorophyll. Cotyledon one, linear, photosynthesizing, dorsiventrally flattened (bifacial). Cotyledon hyperphyll prolonged, assimilating. Hypocotyl internode absent. Coleoptile absent. Collar rhizoids present. Radicula very short. Germination phanerocotylar.

Phytochemistry O-methylated flavonols (kaempferol, quercetin, syringetin), cyanidin, delphinidin, and proanthocyanidins present. Ferulic and diferulic acids present in cell walls. Ellagic acid, alkaloids, saponins, and cyanogenic compounds not found.

Kunth in von Humboldt, Bonpland et Kunth, Nov. Gen. Sp. Plant. 1, ed. 4°: 265. Mai 1816 [‘Pontedereae’], nom. cons.

Pontederiales A. Rich. in C. F. P. von Martius, Consp. Regn. Veg.: 7. Sep-Oct 1835 [‘Pontederaceae’]; Unisemataceae Raf., New Fl. N. Amer. 2: 75. Jul-Dec 1837 [’Unisemides’], nom. illeg.; Heterantheraceae J. Agardh, Theoria Syst. Plant.: 36. Apr-Sep 1858 [’Heteranthereae’]; Pontederianae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992

Distribution Pantropical, with their largest diversity in tropical America, some species in subtropical and warm-temperate regions on both hemispheres.

Fossils A fossilized shoot with roots, which is similar to extant Monochoria or ‘Eichhornia’, was described from the Maastrichtian Deccan Intertrappean Beds in India. Pontederites, which may be assigned to Pontederiaceae, has been found in Eocene layers in the United States, and seeds and leaves of Pontederiaceae occur in Cenozoic layers.

Habit Bisexual, usually perennial (sometimes annual) herbs. Aquatic (sometimes freely floating) or helophytes. Vegetative stems with indeterminate growth.

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vessels in root (in some species ‘Eichhornia’ and Heteranthera also in stem). Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids with simple? pits. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Idioblasts (myriophyllin cells) with condensed tannins present in some species. Laticifers absent. Calciumoxalate raphides present, in Pontederia also cells with styloids or one or two prismatic crystals. Tanniniferous cells (idioblasts) sometimes present in pseudopetioles. Silica bodies absent.

Leaves Alternate (usually distichous, sometimes spiral; rarely verticillate), simple, entire, linear or differentiated into pseudopetiole and pseudolamina (pseudopetiole in ‘Eichhornia’ swollen and air-filled); pseudolamina of younger leaves often enclosing pseudopetiole of older leaves (leaves in Hydrothrix strongly reduced, filiform), with ? ptyxis. Stipules absent; leaf sheath short, open or closed, often as an axillary or median ochrea-like structure, large and enclosing stem, or reduced to a ligule or absent. Colleters present. Venation parallelodromous with converging veins or pinnate-parallel, acrodromous; primary veins longitudinal and connected to transversal secondary veins. Stomata paracytic; adjacent cells with oblique divisions. Cuticular wax crystalloids absent. Mesophyll with calciumoxalate raphides or single prismatic crystals. Leaf margin entire.

Inflorescence Terminal, panicle, thyrse, raceme-, spike- or umbel-like, subtended by two basal foliaceous or sheathing bracts, upper one often a spatha (flowers sometimes solitary; in Hydrothrix enantiomery: two flowers in each pair mirror images, densely inserted inside a spatha and forming a pseudanthium).

Flowers Zygomorphic (sometimes almost actinomorphic; sometimes obliquely zygomorphic), sometimes large. Hypogyny. Tepals usually 3+3 (sometimes three or 2+2), pseudomonocyclic, petaloid (outer tepals in Monochoria almost sepaloid), usually early caducous, more or less connate at base into a tube. Septal nectaries present in ‘Eichhornia’ and Pontederia. Triheterostyly and pollen trimorphism present in species of Pontederia and ‘Eichhornia’; (somatic) enantiostyly with left-hand and right-hand oriented flowers and staminal dimorphism present in Monochoria and Heteranthera.

Androecium Stamens usually 3+3 (sometimes 2+2, three or one, by reduction of outer staminal whorl). Filaments glandular hairy or with moniliform hairs, free, adnate to tepals. Anthers basifixed (sometimes auriculate and seemingly dorsifixed), sometimes versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores). Tapetum amoeboid-periplasmodial or secretory, with ab initio uninucleate or binucleate cells. Staminodia usually absent (sometimes two or three, when fertile stamens fewer than six).

Pollen grains Microsporogenesis successive. Pollen grains usually disulculate (rarely monosulculate or trisulculate), shed as monads, bicellular or tricellular at dispersal. Exine intectate, acolumellate, with pollen surface verrucate, scabrate or finely reticulate.

Gynoecium Pistil composed of three connate carpels (in Pontederia pseudomonomerous, with only median carpel developing and fertile). Ovary superior, unilocular (sometimes due to reduction of two carpels) or trilocular. Style single, simple, glabrous or hairy. Stigma small, capitate or trilobate, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (when ovary unilocular, then seemingly parietal, with intrusive placentae). Ovules one per ovary or usually ten to more than 50 per carpel, anatropous, pendulous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument two cell layers thick. Inner integument ? cell layers thick. Parietal cell formed from archesporial cell but usually not forming a parietal tissue (parietal tissue in Monochoria one cell layer thick). Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development helobial (cell wall formation starting much earlier in small chalazal chamber than in large micropylar chamber). Endosperm haustoria at least in Monochoria two micropylar, developing from each side of chalazal chamber and protruding into chalazal tissue, micropylar chamber subsequently fusing with haustoria. Embryogenesis asterad.

Fruit Usually a loculicidal or irregularly dehiscent capsule (in Pontederia an achene, surrounded by hardening persistent petiole bases, forming an anthocarp).

Seeds Aril absent. Seed coat testal. Exotestal cells longitudinally elongate, box-shaped. Endotestal cells thin, transversally elongate, with prolonged end-walls, with silica crystals? Tegmen crushed, containing a reddish-brown pigment. Operculum present (combined with the micropylar collar?). Perisperm not developed. Endosperm copious, starchy, with simple starch grains. Suspensor little or not developed. Embryo large, straight, well differentiated, Trillium type, chlorophyll? Cotyledon one, linear, bifacial. Hypocotyl internode absent. Cotyledon hyperphyll elongate, assimilating. Coleoptile absent (tubular cotyledon ligule present). Collar rhizoids present. Plumule lateral. Germination phanerocotylar? First leaves of seedling linear.

Cytology n = (7) 8 (9, 13–17, 20, 21, 24, 26, 29, 30, 36, 37, 40, 41) – Polyploidy and aneuploidy occurring.

Phytochemistry O-methylated flavonols, cyanidin, proanthocyanidins (prodelphinidins), alkaloids, cyanogenic compounds, phenylphenalenones, arylphenalenones, and polyamines present. Diferulic and p-coumaric acids in cell walls? Ellagic acid chelidonic acid, and saponins not found.

Systematics ‘Eichhornia’ (7; E. azurea, E. crassipes, E. diversifolia, E. heterosperma, E. natans, E. paniculata, E. paradoxa; southeastern United States?, Mexico, Central America, the West Indies, tropical South America; incl. Heteranthera, Monochoria and Pontederia etc?), Monochoria (6; M. africana, M. cyanea, M. elata, M. hastata, M. korsakowii, M. vaginalis; northeastern Africa to Japan and Austrtalia; in Eichhornia?), Pontederia (6; P. rotundifolia, P. sagittata, P. subovata, P. triflora, P. angustifolia; Canada to Argentina; in Eichhornia?), Heteranthera (9; tropical and subtropical Africa and America; in Eichhornia), Hydrothrix (1; H. gardneri; northeastern Brazil; in Eichhornia?), Eurystemon (1; E. mexicanum; Mexico, northern Central America; in Eichhornia?), Scholleropsis (1; S. lutea; Lake Chad in northern Cameroon, Madagascar; in Eichhornia?).

Pontederiaceae are probably sister-group to Haemodoraceae.

Generic delimitations are unclear. ’Eichhornia’ is paraphyletic and should either include the entire Pontederiaceae or be split. Zosterella is included in Heteranthera.

In Heteranthera, Eichhornia s. str. and Pontederia s. str. the leaves of the axillary shoot entirely enclose the main stem in bud, or the pseudolamina of the young leaf completely enclose the pseudopetiole of the second oldest leaf. This may be a synapomorphy.

Cladogram (simplified) of Pontederiaceae based on DNA sequence data (Kohn & al. 1996). Eichhornia meyeri was identified as sister-group to the remaining Pontederiaceae in the analysis by Ness & al. (2011).

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COMMELINANAE Takht.

COMMELINALES Mirb. ex Bercht. et J. Presl

COMMELINACEAE Mirb.

HAEMODORACEAE R. Br.

HANGUANACEAE Airy Shaw

PHILYDRACEAE Link

PONTEDERIACEAE Kunth