[Iridales+Commelinidae]


IRIDALES Raf.

Rafinesque, Anal. Nat.: 197. Apr-Jun 1815 [’Iridisia’]

Iridanae Doweld, Tent. Syst. Plant. Vasc.: lvi. 23 Dec 2001

Habit Usually bisexual (rarely monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, gynodioecious, or polygamodioecious), usually perennial herbs (sometimes biennial herbs, or evergreen or deciduous trees, shrubs, suffrutices or lianas). Often with bulb or corm rich in polysaccharides. Often xerophytic.

Vegetative anatomy Usually Arum type arbuscular mycorrhiza. Single- or multi-layered velamen often present. Phellogen absent. Secondary lateral growth usually absent (sometimes anomalous). Vessels present only 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 P2c or P2cf types (rarely P2cs type). Nodes usually multilacunar with several leaf traces. Laticifers sometimes present. Silica bodies sometimes present. Mucilage cells and mucilage sacs frequent, with calciumoxalate as raphides, pseudo-raphides, styloids, cubes, crystal sand, or rhomboidal or acicular crystals. Epidermal cells often with crystals.

Trichomes Hairs usually absent (sometimes unicellular or multicellular, uniseriate or multiseriate, stellate or lepidote, rarely dendritic).

Leaves Alternate (spiral or distichous), simple, entire, often linear, usually bifacial (sometimes equitant), with revolute, supervolute, involute, conduplicate, plicate, flat or curved (sometimes explicative) ptyxis, sometimes differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath open or closed, often well developed (sometimes absent). Venation parallelodromous or pinnate-parallelodromous (sometimes acrodromous). Stomata anomocytic, paracytic, tricytic or tetracytic; neighbouring cells usually with oblique or non-oblique divisions. Cuticular wax crystalloids as parallel platelets (Convallaria type, resembling ‘electromagnetic field lines’) or unordered platelets, rodlets or filiform reticulate processes. Dimorphic hypodermal cells sometimes present. Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin usually entire (sometimes serrate).

Inflorescence Terminal, umbel-, raceme-, spike- or head-like, panicle, fascicle, corymb, or thyrsoid, often compound and consisting of bostrychoid or helicoid monochasial partial inflorescences, or racemes or spikes, sometimes subtended by membranous spathae; or flowers solitary. Floral prophylls (bracteoles) sometimes lateral, pairwise or absent.

Flowers Actinomorphic or zygomorphic. Hypogyny or epigyny (rarely half epigyny). Tepals (2–)3(–5)+(2–)3(–5) (rarely 7+7), usually petaloid (sometimes sepaloid), usually more or less connate into infundibuliform or tubular perianth (rarely free). Septal nectaries present, usually infralocular, or absent. Disc usually absent (sometimes nectar-secreting).

Androecium Stamens (2–)3(–6)+(2–)3(–6) (sometimes one or three, rarely two, five, nine to more than 18). Filaments free or more or less connate, often more or less adnate to tepals (rarely to style). Anthers usually dorsifixed or basifixed (sometimes centrifixed), versatile or non-versatile, tetrasporangiate, usually introrse (sometimes latrorse, rarely extrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pore). Tapetum usually secretory (rarely amoeboid-periplasmodial), with binucleate to quadrinucleate cells. Staminodia three or absent (female flowers often with staminodia). Pollinaria present in most Orchidaceae, consisting of pollinia and translator.

Pollen grains Microsporogenesis usually successive (sometimes simultaneous). Pollen grains usually monosulcate (sometimes monosulcoidate or inaperturate, rarely disulcate, disulculate, trisulcate, trichotomosulcate, zonosulcate, zonosulculate, dizonosulculate, spiraperturate, mono- to tetraporate, ulcerate, or foraminate), usually shed as monads (rarely tetrads), bicellular at dispersal. Exine tectate or semitectate (rarely intectate), with columellate infratectum, reticulate, rugulate, foveolate, echinate, verrucate, spinulate, or smooth (sometimes absent).

Gynoecium Pistil composed of (two or) three (to seven) connate carpels. Ovary superior or inferior (rarely semi-inferior), (unilocular to) trilocular (to quinquelocular). Style single, simple, with stylar canal (rarely trilobate). Stigma capitate, punctate or slightly to deeply trilobate, papillate or non-papillate, Dry or Wet type. Pistillodium usually absent (male flowers often with pistillodium).

Ovules Placentation usually axile (sometimes parietal, rarely basal or apical). Ovules one to more than 50 per carpel (sometimes hundreds to several thousands, rarely to four million per ovary), anatropous or campylotropous (sometimes semicampylotropous, hemianatropous or orthotropous), ascending or horizontal, apotropous or epitropous, usually bitegmic (sometimes unitegmic, rarely ategmic), usually crassinucellar (sometimes tenuinucellar, sometimes pseudocrassinucellar or pseudotenuinucellar). Micropyle endostomal or bistomal (rarely exostomal). Funicular obturator sometimes present. Parietal cell formed from archesporial cell or absent. Periclinal cell divisions often occurring in megasporangial epidermis. Nucellar cap sometimes present. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, rarely tetrasporous, Scilla, Adoxa, Fritillaria, or Drusa type). Synergids sometimes with a filiform apparatus. Antipodal cells usually persistent, sometimes proliferating. Endosperm development usually ab initio helobial (sometimes nuclear). Endosperm haustoria chalazal or absent. Embryogenesis asterad or onagrad (sometimes caryophyllad, chenopodiad or solanad).

Fruit Usually a loculicidal capsule (rarely septicidal, irregularly dehiscent or indehiscent) or a berry (sometimes a nut, rarely a drupe, a samara or a schizocarp).

Seeds Aril usually absent. Strophiolus rarely present. Caruncular, chalazal or raphal elaiosome occasionally present. Seed coat usually exotestal, sometimes fleshy or winged, sometimes with caruncular elaiosome at chalazal end. Exotesta usually with thin phytomelan layer on epidermal cell walls. Mesotesta and endotesta often collapsed. Tegmen usually collapsed, sometimes well developed. Perisperm usually not developed (sometimes well developed, with lipids and proteins). Endosperm copious, with oils, aleurone or hemicellulose (sometimes rich in water and/or starch), sometimes thin-walled. Embryo straight or slightly curved, little differentiated, with or without chlorophyll. Cotyledon one, sometimes photosynthesizing. Cotyledon hyperphyll elongate, dorsiventrally flattened and assimilating, or compact and not assimilating. Hypocotyl internode short or absent. Coleoptile absent. Radicula well developed, contractile. Germination phanerocotylar or cryptocotylar.

Cytology n = 2–23

DNA Mitochondrial genes rpl2 and sdh3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, isorhamnetin), flavone-C-glycosides, lanaroflavone (biflavonoid), isoflavones, amentoflavone (biflavone), homoisoflavanones, cyanidin, chalcones, 6-hydroxyapigenin methyl ethers, tannins, phenols, norbelladine alkaloids (toxic tyrosine derivatives, e.g. crinine, belladine, galanthanine, haemanthanine, homolyrine, lycorenine, lycorine, pancratiostatine, and tazettine), polyhydroxyalkaloids, cholestane glycosides (cardiotoxic bufodienolides, cardenolide glycosides and spirostanol glycosides), phenolic glycosides, steroidal saponins and sapogenins (e.g. agapanthagenin, yuccagenin, diosgenin, hecogenin, neotigogenin, sarsasapogenin, smilagenin, tigogenin, yamogenin, aigogenin, markogenin diglycoside, timosaponins, anemarsaponins, gitogenin, aspidirin), cyanogenic compounds (e.g. cyanogenic glycosides), anthrones (e.g. anthrone-C-glycoside [aloin, barbaloin] in leaves), tetrahydroanthracenones (chrysophanol, 10,7’-bichrysophanol, knipholone, asphodeline and other lipophilic anthranoid aglucones), naphthoquinones (imbricatonol, plumbagin, stypandrone), quinonoid pigments, polyacetate-derived arthroquinones, nepodin, dianellidin, stypandrol, dianellidone, magniferin (glycosylic xanthone), chromones, phenylpyrones, phenolic amines, acetidine carbonic acid, chelidonic acid, ascorbic acid, tuliposides, allyl sulfides, allyl disulfides, propyl sulfides, vinyl disulfides, alliin, propionaldehyde, propionthiol, hydroxycinnamic acid, eicosanyl arachidate, crocein, phytosterols, non-protein amino acids (tricine etc.), meta-carboxysubstituted aromatic amino- and γ-glutamic peptides, acaroid resins, and stem fructans present. Ellagic acid not found.

Systematics Iridales are sister-group to Commelinidae. Orchidaceae seem to be sister to the remaining Iridales. Stevens (2001 onwards) has suggested potential synapomorphies for some of the main clades in Iridales.

The clade [[Boryaceae-Hypoxidaceae]+[Ixioliriaceae+Tecophilaeaceae]+[Doryanthaceae-Ruscaceae]] is characterized by, e.g.: cuticular wax crystalloids as parallel platelets; tepals often connate; stamens often inserted on a perigonal tube; seed with well developed exotesta and phytomelan layer; and presence of stem fructans.

The first of these main clades is [Boryaceae+[Blandfordiaceae+[Lanariaceae+[Asteliaceae+ Hypoxidaceae]]]], which has the following potential synapomorphies: septal nectaries external; hypostase often present in ovule; megagametophyte often with chalazal constriction; and antipodal cells usually persistent. The clade [Blandfordiaceae+[Lanariaceae+[Asteliaceae+Hypoxidaceae]]] is characterized by ovule usually with parietal cell and nucellar cap. The [Lanariaceae+ [Asteliaceae+Hypoxidaceae]] has: multicellular and often branched hairs; paracytic stomata; pseudolamina with distinct midrib; often internal septal nectaries; and bistomal micropyle. Finally, potential synapomorphies of the clade [Asteliaceae+Hypoxidaceae] include presence of mucilage canals; thin-walled endosperm; cotyledon not photosynthesizing; an elongate ligule; and presence of flavonols (Stevens 2001 onwards).

Ixioliriaceae and Tecophilaeaceae have the following characteristics in common (Stevens 2001 onwards): cormose stem; spiral leaves; outer tepals mucronate to aristate; short perigonal tube; and stamens arising from the mouth of the perianth tube.

The second main clade is [Iridaceae+[Xeronemataceae-Ruscaceae]], and this has the potential synapomorphies (Stevens 2001 onwards): often absence (loss) of Arabidopsis type telomeres; human-type telomere ([TTAGGG]n type telomere) dominating; often glucomannans as reserve carbohydrates; and often secondary growth (from meristem producing tissue inwards; separate vascular bundles in this tissue surrounded by ground parenchyma). The clade [Xeronemataceae+[Hemerocallidaceae-Ruscaceae]] has lost the mitochondrial gene rpl2. Characteristic features of the clade [[Hemerocallidaceae+[Xanthorrhoeaceae+Asphodelaceae]]+ [Alliaceae-Ruscaceae]] includes: pedicels articulated; septal nectaries infralocular; hypogyny; microsporogenesis successive; and parietal tissue two or three cell layers thick. The clade [Hemerocallidaceae+[Xanthorrhoeaceae+Asphodelaceae]] has: styloids; cymose inflorescence branches; outer integument more than three cell layers thick; hypostase; non-photosynthesizing cotyledon; and anthraquinones. [Xanthorrhoeaceae+Asphodelaceae] is characterized by: secondary thickening and lateral growth sometimes present; stamens not adnate to tepals; and angular seeds.

The monophyletic group [[Agapanthaceae+[Amaryllidaceae+Alliaceae]]+[Aphyllanthaceae-Ruscaceae]] has the potential synapomorphy of successive microsporogenesis. The [Agapanthaceae+[Amaryllidaceae+Alliaceae]] clade has the common features (Stevens 2001 onwards): presence of laticifers and mucilage cells; leaves distichous; inflorescence scapose, umbel-like cymose, with scarious spatha, with two (or more, external) inflorescence bracts; pedicels not articulated; stigma usually Dry type; parietal tissue absent; presence of flavonols and saponins; and lectins binding mannose (in Agapanthus?).

The clade [[[Aphyllanthaceae+[Themidaceae+Hyacinthaceae]+[Anemarrhenaceae-Behniaceae]]+[Laxmanniaceae+[Asparagaceae+Ruscaceae]]] has racemose inflorescence. Themidaceae and Hyacinthaceae share the features: spiral leaves; inflorescence scapose, with bracteate pedicels; presence of raphides in carpel wall; anatropous ovules; non-photosyntesizing cotyledon; and presence of steroidal saponins. In [Agavaceae+[Behniaceae+[Anthericaceae+ Herreriaceae]]] are nucellar cap and hypostase present, whereas articulated pedicels, capsular fruit, a thick-walled, pitted and hemicellulosic helobial endosperm, and presence of steroidal saponins are characteristic features of the clade [Laxmanniaceae+[Asparagaceae+Ruscaceae]].


Phylogeny (maximum likelihood bootstrap consensus tree) of Iridales based on DNA sequence data (Wurdack & Dorr 2009).

AGAPANTHACEAE F. Voigt

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Voigt, Gesch. Pflanzen-Reichs 2: 440. 1850 [’Agapantheae’]

Genera/species 1/6–9

Distribution Southern and southeastern South Africa.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome tuberous.

Vegetative anatomy Roots fleshy, contractile, with multiple velamen. Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements usually with scalariform (sometimes simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Laticifers absent? Calciumoxalate raphides present.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, linear, with flat ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids? Mesophyll with calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, umbel-like, consisting of condensed helicoid monochasia, subtended by two membranous caducous bracts (spathae) connate on one side, which enclosed floral buds. Peduncle (scape) long and stout, terete or somewhat compressed. Floral bracts and prophylls filiform, persistent.

Flowers Zygomorphic, large. Pedicel not articulated. Hypogyny. Tepals 3+3, petaloid, fleshy, with abaxial median ridge and adaxial median furrow, more or less connate at base. Septal nectaries infralocular. Disc absent.

Androecium Stamens 3+3. Filaments free from each other, adnate at base to tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

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

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple, narrow, curved, with stylar canal. Stigma small, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules numerous per carpel, semi-campylotropous, apotropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal cell formed from archesporial cell. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit A loculicidal capsule with persistent style.

Seeds Aril absent. Testa winged at funicular end. Seed coat exotestal. Exotesta with thin phytomelan layer on epidermal cell walls. Mesotesta, endotesta and tegmen collapsed. Perisperm not developed. Endosperm copious, with oil, aleuronic, starch and hemicellulose. Embryo large, well developed, without chlorophyll. Cotyledon one. Germination cryptocotylar.

Cytology n = (14) 15 (16) – Chromosomes 4–9 µm long.

DNA The mitochondrial gene rpl2 is absent.

Phytochemistry Insufficiently known. Steroidal saponins (e.g. agapanthagenin and yuccagenin) present. Flavonols, allyl sulphides, alliin etc. not found.

Use Ornamental plants.

Systematics Agapanthus (6–9; southern and southeastern South Africa from Western Cape northwards to just south of Limpopo River).

Agapanthus is sister to [Alliaceae+Amaryllidaceae].

AGAVACEAE Dumort.

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Dumortier, Anal. Fam. Plant.: 57, 58. 1829 [‘Agavineae’], nom. cons.

Funkiaceae Horan., Prim. Lin. Syst. Nat.: 52. 2 Nov 1834 [’Funkiaceae s. Hemerocallideae’], nom. illeg.; Yuccaceae J. Agardh, Theoria Syst. Plant.: 8. Apr-Sep 1858; Agavales Hutch., Fam. Fl. Pl., Monocot. 2: 14, 149. 20 Jul 1934; Hesperocallidaceae Traub in Plant Life 28: 131. 22 Feb 1972 [’Hesperocallaceae’]; Hostaceae B. Mathew in Kew Bull. 43: 302. Jun 1988; Chlorogalaceae (Baker) Doweld et Reveal in Reveal in Bot. Rev. (Lancaster) 71: 53. 20 Mai 2005

Genera/species 10/350–375

Distribution East Asia, Central and southwestern North America and southwards to tropical and subtropical South America and the West Indies, with their largest diversity in Mexico.

Fossils Protoyucca shadishii, resembling extant Yucca, is represented by fossilized stems from the mid-Miocene of Nevada.

Habit Usually bisexual (rarely andromonoecious, gynomonoecious or dioecious), usually perennial herbs, sometimes giant herbs (Agave, Furcraea, etc.; some species of Yucca are arborescent). Rhizome often tuberous; in Hesperocallis a bulb with tunica and onion-like smell. Many species are xerophytic.

Vegetative anatomy Roots fibrous or fleshy, in Agave with multilayered velamen; root endodermis in Yucca thick. Phellogen? Primary vascular tissue as scattered bundles (atactostele). Secondary lateral growth usually absent (Agave, Furcraea and Yucca have anomalous secondary lateral growth). Vessels present in roots. Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes multilacunar with ? leaf traces. Mucilage cells abundant. Silica bodies absent. Calciumoxalate as pseudo-raphides (in Agave hexagonal in cross-section), raphides, styloids, and prismatic or acicular crystals.

Trichomes Hairs usually absent (sometimes unicellular or multicellular, uniseriate).

Leaves Alternate (spiral), simple, entire, often linear, sometimes succulent (in Hosta sometimes differentiated into pseudopetiole and pseudolamina), with ? ptyxis. Stipules absent; leaf sheath well developed. Adaxial vascular bundles inverted. Venation parallelodromous (in Hosta sometimes pinnate-parallelodromous); primary veins in Hosta usually distinct, connected by transversal secondary veins. Stomata anomocytic, paracytic, tricytic or tetracytic, sunken, in Agave, Beschorneria, Furcraea, ‘Prochnyanthes’, and Yucca surrounded by complex spaces. Cuticle in Agave, Beschorneria, Furcraea, ’Prochnyanthes’, and Yucca thick. Cuticular wax crystalloids usually as parallel platelets (Convallaria type; in Hosta as non-orientated tubuli, chemically dominated by nonacosan-10-ol). Mesophyll with calciumoxalate as raphides or single prismatic crystals, without mucilaginous idioblasts. Bundles of strongly lignified fibres present near leaf surface; particularly long and strong fibres present in species of Agave, Hesperaloe and Yucca. Vascular bundles in Agave and Yucca with acicular calciumoxalate crystals (sometimes with styloids or suberized cells containing pseudo-raphides). Leaf margin usually entire (sometimes serrate, with tiny translucent teeth, in Agave and Furcraea with large hard teeth), in some species filiferous and separated by layers of lignified cells and abscission cells. Leaf apex often acute (in Agave, Hesperaloe and Yucca with a hard sclerenchymatous terminal spine).

Inflorescence Terminal or axillary, panicle, raceme- or spike-like, with one- to many-flowered partial inflorescences. Bracts caducous or persistent.

Flowers Actinomorphic or zygomorphic, usually large. Pedicel usually not articulated and without pericladium (in Chlorogalum, Hesperocallis and Schoenolirion articulated). Hypogyny to epigyny. Tepals 3+3 (median outer tepal adaxial), petaloid, caducous or marcescent, usually connate into a tube, often connate at base (sometimes free). Septal nectaries present or absent. Disc absent.

Androecium Stamens 3+3 (sometimes three fertile and three staminodial). Filaments narrow to wide and thick, free from each other, usually adnate to tepals (epitepalous) (filaments in some species of Hosta inserted on ovary; filaments in some species of Yucca hairy). Anthers dorsifixed, versatile, tetrasporangiate, usually introrse, longicidal (dehiscing by longitudinal slits); connective in Hosta cucullate and widening around filament apex (filaments inserted in a pit on connective). Tapetum secretory, with binucleate to quadrinucleate cells. Staminodia three or absent.

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (rarely disulculate), usually shed as monads (rarely tetrads), bicellular at dispersal. Exine tectate or semitectate, with ? infratectum, usually reticulate (in Hosta usually rugulate or verrucate).

Gynoecium Pistil composed of three connate carpels. Ovary superior to inferior, trilocular. Style usually single, simple (stylodia in most species of Yucca three, free), usually narrow (in Beschorneria, Furcraea and Yucca thick; in Hesperocallis persistent; in Camassia with three canals). Stigma capitate, punctate or trilobate (stigmas in most species of Yucca three), papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules six to more than 50 per carpel, anatropous, bitegmic, usually crassinucellar (sometimes tenuinucellar). Micropyle ?-stomal. Outer integument (four to) nine to 14 cell layers thick. Inner integument two cell layers thick. Obturator present in Beschorneria. Parietal cell formed from archesporial cell (also in Hosta and Hesperocallis). Hypostase present (also in Hosta). Parietal tissue one or two cell layers thick or absent. Nucellar cap present, sometimes two cell layers thick. Megagametophyte monosporous, Polygonum type (micropylar megaspore cell dividing in Furcraea). Synergids sometimes with a filiform apparatus. Antipodal cells persistent. Chalazal part of megagametophyte possibly with haustorial function. Endosperm development usually helobial (also in Hesperocallis and Hosta; in Camassia and Furcraea nuclear). Endosperm haustoria absent. Embryogenesis onagrad or caryophyllad (Yucca).

Fruit Usually a loculicidal capsule (in Yucca a septicidal and/or loculicidal capsule, a dry indehiscent fruit or a berry).

Seeds Aril absent. Seeds usually flat, sometimes winged. Seed coat testal. Exotesta with a thin layer of phytomelan on epidermal cell walls. Mesotesta and endotesta present in, e.g., Agave. Tegmen collapsed. Perisperm in Agave and Yucca conspicuous, with proteins and lipids (starch absent). Endosperm usually copious (in Yucca ruminate), with oil, aleurone, protein and hemicellulose (starch absent); endosperm with usually thick pitted cell walls (in Hosta with thin cell walls). Embryo large, straight or slightly curved, without chlorophyll. Cotyledon one, photosynthesizing (in Hosta little differentiated, not photosynthesizing). Cotyledon hyperphyll usually elongate, assimilating (in Hosta compact, not photosynthesizing). Hypocotyl internode up to 4 mm long. Coleoptile absent. Radicula persistent (Agave, Yucca) or ephemeral, often branched. Collar rhizoids usually present. Germination in Hosta cryptocotylar.

Cytology n = 30; n = 24 (Hesperocallis) – Polyploidy and aneuploidy occurring in, e.g., Agave and Hosta. Chromosomes 0,4–10 µm long. Karyotype bimodal, with four (Hosta), five (Agave, Yucca), or five or six (Hesperocallis) long, and c. 25 short chromosomes (in Hesperocallis 18 or 19 short chromosomes; in Hosta also two medium-sized chromosomes). Adventitious polyembryony present in Hosta (at least in H. ventricosa).

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols and other flavonoids (kaempferol xyloside, kaempferol-3-glycoside, kaempferol-3-rutinoside, etc.), steroidal saponins (diosgenin, hecogenin, neotigogenin, sarsasapogenin, smilagenin, tigogenin, yamogenin and nearly 30 other sapogenins, especially in seeds), chelidonic and ascorbic acid, non-protein amino acids, and fructans present. Homoisoflavanones present in Chlorogaleae. Ellagic acid, tannins, proanthocyanidins, alkaloids, and cyanogenic compounds not found.

Use Ornamental plants, textile plants (fibres from Agave sisalana and A. fourcroydes), soap (Yucca), medicinal plants, stimulants and beverages (Agave tequilana).

Systematics Hesperocallis (1; H. undulata; southern California, western Arizona); Hosta (c 40; China, the Korean Peninsula, Japan); Schoenolirion (3; California, southeastern United States), Hastingsia (4; western North America), Camassia (6; North America), Chlorogalum (5; California); Hesperoyucca (3; southwestern United States, northwestern Mexico), Hesperaloe (5; southwestern United States, northwestern Mexico), Yucca (c 50; central United States, California, Mexico, Central America, the West Indies), Furcraea (c 20; tropical America), Beschorneria (7; central and southern Mexico), Agave (c 240; southern United States, Mexico, Central America, the West Indies, Venezuela).

Agavaceae are sister-group to the clade [Anthericaceae+[Behniaceae+Herreriaceae]].

Hosta is sister to the remaining Agavaceae in Bogler & Simpson (1996). On the other hand, Hesperocallis is sister to Agavaceae in analyses by Seberg & al. (2012). Hesperocallis undulata and Hosta plantaginea have pollen grains with similar unibaculate muri.

Chlorogaleae Baker in Bot. J. Linn. Soc. London 13: 215. 4 Dec 1872 – comprising Camassia, Chlorogalum, Hastingsia and Schoenolirion – are successive sister-group to Agavaceae minus Hesperocallis and Hosta. They have rhexigenetic lacunae like Scilloideae (Hyacinthaceae) and at least Camassia has tepals with a simple leaf trace. In a Bayesian analysis of molecular data (Kim & al. 2010), Camassia was recovered with very high support as sister to the clade [Agavaceae+[Anthericaceae+[Behniaceae+Herreriaceae]]]. Unfortunately, Chlorogalum, Hastingsia or Schoenolirion were not included in that study. Awaiting analyses of all four genera, they will be kept in Agavaceae.

Cladogram of Agavaceae based on DNA sequence data (Bogler & Simpson 1996). Hesperocallis is sister to the remaining Agavaceae in analyses by Seberg & al. (2012).

Strict consensus tree (simplified) of Agavaceae based on DNA sequence data (Bogler & al. 2006).

ALLIACEAE Batsch ex Borkh.

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Borkhausen, Bot. Wörterb. 1: 15. 1797, nom. cons.

Alliales Bercht. et J. Presl, Přir. Rostlin: 267. Jan-Apr 1820 [’Alliaceae’]; Gilliesiaceae Lindl. in Bot. Reg. 12: ad t. 992. 1 Jul 1826 [’Gilliesieae’]; Alliineae Link, Handbuch 1: 152, 820. 4-11 Jul 1829 [‘Alliaceae’]; Gilliesiales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 7. Sep-Oct 1835 [‘Gilliesieae’]; Cepaceae Salisb., Gen. Plant.: 88. 15-31 Mai 1866 [’Cepaeeae’]; Tulbaghiaceae Salisb., Gen. Plant.: 87. 15-31 Mai 1866 [’Tulbagheae’]; Milulaceae Traub in Plant Life 28: 131. 22 Feb 1972

Genera/species 12/850–890

Distribution Temperate and subtropical regions in the Northern Hemisphere, South Africa, Sri Lanka, Mexico.

Fossils Unknown.

Habit Bisexual, perennial or biennial herbs. Usually with a bulb surrounded by membranous scales (rarely a bulb-like corm; in Tulbaghia a tuberous rhizome). Often with a strong characteristic odour (‘onion smell’). New bulb developing in axile of uppermost leaf.

Vegetative anatomy Roots of young bulb often contractile. Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform and/or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Laticifers present in at least Allium, Nothoscordum, Tristama, and Tulbaghia. Mucilage cells abundant. Calciumoxalate raphides present in some genera, absent from at least Allium, Milula, and Tulbaghia; calciumoxalate as single crystals, styloids or other configurations present in Allium and Nothoscordum.

Trichomes Hairs unicellular or absent.

Leaves Alternate (spiral or distichous), simple, entire, often linear or filiform, often fleshy, often angular in transverse section or terete, with ? ptyxis. Stipules absent; leaf sheath closed, long, in Allium etc. with short ligule. Venation parallelodromous; vascular bundles in many species of Allium (especially with cylindrical leaves) arranged in a cylinder with phloem adjacent to epidermis. Stomata anomocytic. Cuticular wax crystalloids as platelets without particular orientation (in several species of Allium), as tubuli (chemically dominated by nonacosan-10-ol) or as filiform reticulate processes (in Ipheion uniflorum and Nothoscordum bivalve). Mesophyll often with calciumoxalate raphides. Leaf margin entire.

Inflorescence Usually terminal (rarely lateral), umbel-like (in one species of Milula spike-like), consisting of one or several condensed helicoid monochasia (rarely a single flower). Inflorescence subtended by usually two (sometimes one or more than two) membranous, usually free (sometimes connate) bracts (spathae), enclosing floral buds and in some species caducous prior to anthesis. Peduncle (scape) usually long and terete (occasionally angular or somewhat compressed in transverse section). Floral bracts and prophylls usually absent. Flowers in some species of Allium often replaced by bulbils.

Flowers Usually actinomorphic (in Gilliesia, Miersia and Solaria zygomorphic). Usually hypogyny (in Allium siculum and A. tripedale almost half epigyny). Tepals usually 3+3 (rarely three or 3+2), petaloid, free or more or less connate; usually with appendages of various shape between tepals or stamens, often as corona. Septal nectaries infralocular (in Allium with orifice close to ovary base; in Nothoscordum with orifice near style). Disc absent.

Androecium Stamens usually 3+3 (rarely two or three stamens, or two fertile stamens and three staminodia, or three fertile stamens and two staminodia). Filaments free or connate, often adnate to tepals (in some species with apical, dorsal or lateral appendages). Anthers basifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate (Allium) or binucleate to quadrinucleate (Tulbaghia) cells. Staminodia usually absent (rarely two or three).

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary usually superior (rarely semi-inferior), trilocular. Style single, simple, terminal or (in Allium and Milula) gynobasic, without stylar canal (style solid). Stigma capitate or trilobate, papillate (with unicellular papillae), usually Dry (in Leucocoryne Wet) type. Pistillodium absent.

Ovules Placentation axile. Ovules two to numerous per carpel, anatropous or campylotropous, apotropous?, bitegmic, crassinucellar or tenuinucellar. Micropyle usually endostomal? (sometimes bistomal). Outer integument ? cell layers thick. Inner integument usually two or three (in Dichelostemma five to seven) cell layers thick. Obturator present (funicular?). Parietal cell not formed (parietal tissue absent). Hypostase present. Nucellar cap usually formed by periclinal cell divisions of megasporangial epidermis. Megagametophyte disporous, Allium type (Allium, Leucocoryne), or monosporous, Polygonum type (Nothoscordum). Endosperm development cellular or nuclear (Allium), or helobial (Nothoscordum, Tulbaghia). Endosperm haustoria? Embryogenesis onagrad or asterad.

Fruit A loculicidal capsule.

Seeds Aril absent. Seed coat exotestal. Exotesta with often coarse phytomelan layer on epidermal cell walls. Mesotesta, endotesta and tegmen usually compressed or collapsed. Perisperm not developed. Endosperm copious, with oil and aleurone (hemicellulose?, starch absent); endosperm cells usually with pitted walls. Embryo short or long, usually curved (in Tulbaghia straight), without chlorophyll. Cotyledon one, usually elongate, usually photosynthesizing. Hypocotyl internode absent. Coleoptile absent. Germination cryptocotylar.

Cytology n = 4–15 – Chromosomes 2–20 µm long. Polyploidy and adventitious embryony (seed formed from megasporangium) occurring (rarely apospory).Minisatellite telomeres are absent in Allium.

DNA Plastid gene infA lost/defunct (Allium). Mitochondrial gene rpl2 lost. Mitochondrial ribosomal protein genes and succinate dehydrogenase genes in Allium transferred to nuclear genome.

Phytochemistry Flavonoids (quercetin etc., especially in bulb scales), steroidal saponins (with, e.g., aigogenin as sapogenin), chelidonic acid, cysteine derived sulphur compounds (strongly smelling onion oils) in the form of, e.g., allyl disulfides, propyl sulfides, vinyl disulfides, propionaldehyde, and propionthiol present in Allium, Milula, Tulbaghia, Gilliesia, Ipheion, Leucocoryne, and Tristagma; in wounded tissues alliin, an S-substituted cystein derived amino acid, converted into water-soluble allicin, pyruvic acid and ammonia, a reaction catalyzed by alliinase. The strongly smelling compound propantial-S-oxide (in, e.g., Allium cepa, although nearly absent from Allium ampeloprasum and A. sativum) is formed from propenylic sulphuric acid and liberated from crushed cells. Ellagic acid, proanthocyanidins and alkaloids not found. Carbohydrates stored in bulb scales as galactan, raffinose, and fructans in, e.g., Allium; in Nothoscordum, Ipheion, and Tulbaghia as starch. Lectines binding mannose.

Use Ornamental plants, vegetables and spices (Allium spp.).

Systematics Alliaceae are sister-group to Amaryllidaceae.

Allioideae Herb., Amaryllidaceae: 48. late Apr 1837 [’Allieae’]

1/750–780. Allium (750–780; temperate regions on the Northern Hemisphere, northern, northeastern and southern Africa, southern Asia southwards to Sri Lanka, Mexico, with their highest diversity in the Mediterranean area, Central Asia and southwestern North America). – Bulbs without starch. Root vessel elements often with simple perforation plates. Leaves sometimes unifacial. Seemingly bifacial leaves in at least some species of Allium with inverted vascular bundles along adaxial surface and bundles with normal orientation along abaxial surface. Tepals usually connate at base. Median outer tepal adaxial. Corona absent. Filaments often winged, connate at base, adnate to inner tepals. Tapetum with uninucleate cells. Ovary sometimes semi-inferior, sometimes with paired projections. Style entire, gynobasic. Ovules two, epitropous, to 14 per carpel. Endosperm development often? cellular. Caruncle sometimes present. Embryo long, curved. n = (7) 8 (9). Numerous transfers of ribosomal protein and succinate dehydrogenase genes from mitochondrial to nuclear genome. Minisatellite telomeres lost. – Allium is sister to [Tulbaghioideae+ Gilliesioideae] (Fay & Chase 1996).

[Tulbaghioideae+Gilliesioideae]

Bulbs with starch. Endosperm development helobial.

Tulbaghioideae (Endl. ex Meisner) M. F. Fay et M. W. Chase in Taxon 45: 448. 13 Aug 1996

1/22. Tulbaghia (22; southern tropical and southern Africa). – Often with rhizome. Leaf sheath short. Floral bracts present. Inner tepals largely connate, often with connate lobes. Corona well developed. Filaments adnate to tepals and/or corona lobes. Ovules two or more per carpel. Seeds flattened. n = 6.

Gilliesioideae Arn., Botany: 134. 9 Mar 1832 [‘Gillesieae’]

c 12/75–85. Zoellnerallium (2; the Andes in central Chile and Argentina), Nothoscordum (20–25; southern and eastern United States to South America), Beauverdia (4; southern Brazil, Uruguay, Argentina), Ipheion (3; Mexico to Chile), Tristagma (17; Chile, Patagonia; incl. Garaventia?), Schickendantziella (1; S. trichosepala; Bolivia, Argentina), Speea (2; Chile), Leucocoryne (15–20; Chile; incl. Erinna?), Trichlora (2; Peru), Solaria (5; southern Chile, southern Argentina), Miersia (3; Bolivia, Chile), Gilliesia (5–6; Chile). – Southern and eastern United States, Mexico, Central America, South America. Flowers in Gilliesia and Miersia very strongly zygomorphic. Tepals in Schickendantziella and Trichlora three, caudate. Corona present or absent. Stamens sometimes two or three (in Gilliesia two). Filaments sometimes connate and adnate to tepals. Anthers sometimes extrorse. Staminodia sometimes present. Style in Nothoscordum entire. Ovules two to numerous per carpel. Embryo short. n = 4 or more. – Zoellnerallium is sister to Nothoscordum and Beauverdia is sister-group to those two (Sassone & al. 2014). Ipheion is sister to Tristagma.

AMARYLLIDACEAE J. St.-Hil.

( Back to Iridales )

Saint-Hilaire, Expos. Fam. Nat. 1: 134. Feb-Apr 1805 [‘Amaryllideae’], nom. cons.

Narcissaceae Juss., Gen. Plant.: 54. 4 Aug 1789 [’Narcissi’]; Leucojaceae Batsch ex Borkh., Bot. Wörterb. 1: 372. 1797; Crinaceae Vest, Anleit. Stud. Bot.: 267, 283. 1818 [‘Crinoideae’]; Amaryllidales Link, Handbuch 1: 193. 4-11 Jul 1829 [’Amarillydeae’]; Narcissales Dumort., Anal. Fam. Plant.: 57. 1829 [‘Narcissarieae’]; Brunsvigiaceae Horan., Prim. Lin. Syst. Nat.: 51. 2 Nov 1834 [‘Brunswigiaceae’]; Crinopsida Horan., Prim. Lin. Syst. Nat.: 51. 2 Nov 1834 [’Crinoideae’]; Galanthaceae G. Mey., Chloris Han.: 510, 560. Jul-Aug 1836 [‘Galantheae’]; Gethyllidaceae Raf., Fl. Tellur. 4: 19. med 1838 [‘Gethylides’]; Amaryllidineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1528. 1846 [‘Amarylleae’]; Narcissineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1528, 1533. 1846 [‘Narcisseae’]; Pancratiaceae Horan., Char. Ess. Fam.: 50. 30 Jun 1847 [‘Pancratiaceae. nob. (Amaryllideae)’]; Cyrtanthaceae Salisb., Gen. Plant.: 138. 15-31 Mai 1866 [‘Cyrtantheae’]; Haemanthaceae Salisb., Gen. Plant.: 129. 15-31 Mai 1866 [‘Haemantheae’]; Oporanthaceae Salisb., Gen. Plant.: 97. 15-31 Mai 1866 [‘Oporantheae’]; Strumariaceae Salisb., Gen. Plant.: 126. 15-31 Mai 1866 [’Strumareae’]; Zephyranthaceae Salisb., Gen. Plant.: 133. 15-31 Mai 1866 [’Zephyrantheae’]

Genera/species 55–57/790–815

Distribution Tropical and subtropical regions on both hemispheres northwards to Western Europe and eastwards to East Asia, with their largest diversity in the Mediterranean, southern Africa and South America.

Fossils Unknown.

Habit Bisexual, perennial herbs. Usually with a bulb rich in polysaccharides (in Clivia, Cryptostephanus and Scadoxus corm or rhizome). Rarely epiphytic or aquatic.

Vegetative anatomy Roots contractile, perennial or fibrous, ephemeral; root velamen two- to four-layered; root exodermis one-layered, with alternating long and short cells. Bulb with tunica. Phellogen absent. Lacunae sometimes formed by degradation of parenchyma. 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 P2cc type (with cuneate protein crystals), P2cco type (with one to three orthogonal and many cuneate protein crystals) (in Worsleya P2ccf type, with cuneate protein crystals and peripheral protein filaments; in Rauhia P2ccof type, with one to three orthogonal and many cuneate protein crystals and peripheral protein filaments). Nodes? Mucilage cells and mucilage sacs frequent, with calciumoxalate raphides and rhomboedric crystals. Epidermal cells often with calcium oxalate crystals.

Trichomes Hairs usually absent (rarely unicellular or multicellular, uniseriate or multiseriate, or aggregated).

Leaves Alternate (usually distichous), simple, entire, often linear, usually bifacial (rarely unifacial), with revolute, involute or flat (sometimes explicative) ptyxis, sometimes differentiated into pseudopetiole and pseudolamina. Stipules absent; leaf sheath sometimes well developed. Venation parallelodromous, sometimes with inverted vascular bundles. Stomata anomocytic, sometimes sunken. Cuticular wax crystalloids as non-orientated rodlets or platelets (similar to those in Asphodelaceae and Hyacinthaceae). Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, umbel- or head-like, consisting of reduced helicoid partial inflorescences (flowers rarely solitary), usually surrounded by (one or) two (to eight) often membranous free or connate equitant or obvolute bracts (spathae), usually marcescent (absent in Leptochiton). Peduncle (scape) often long, sometimes subterranean, in Amaryllideae with a cylinder of sclerenchyma between cortex and central vascular tissue.

Flowers Zygomorphic or (secondarily) actinomorphic, large (some species of Phaedranassa with split zygomorphy, androecial members appearing through an abaxial slit in perianth tube). Pedicel not articulated. Epigyny. Tepals 3+3, petaloid, with median outer tepal adaxial, usually connate in lower part (rarely free) and infundibuliform to tubular. In some clades a corona (paraperigone) present as non-vascularized outgrowths from filaments (in, e.g., Hymenocallis) or vascularized appendages from tepals and forming a tube (e.g. Narcissus), sometimes small and consisting of a narrow border or a ring of scales or fringes in perianth orifice. Septal nectaries usually infralocular (in, e.g., Galanthus tepal nectaries present on distal part of inner tepals; nectariferous glands, associated with septal nectaries, present on filament bases in some species of Eucrosia and Hessea). Disc usually absent (in Galanthus and Leucojum present, nectar-secreting).

Androecium Stamens usually 3+3 (in Zephyra three; in some species of Griffinia five; in Gethyllis 9–18 or more). Filaments free or more or less connate at base, often adnate to perianth tube (in Carpolyza and Strumaria adnate to style), often with prolonged lobed outgrowths forming a corona (e.g. in Hymenocallis). Anthers usually dorsifixed (sometimes centrifixed or basifixed), usually versatile, tetrasporangiate, usually introrse (rarely latrorse), usually longicidal (dehiscing by longitudinal slits; in Galanthus and Leucojum poricidal, dehiscing by an apical pore). Tapetum usually secretory (rarely amoeboid-periplasmodial), with usually binucleate (sometimes uninucleate) cells. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (rarely disulculate), shed as monads, bicellular at dispersal. Exine usually semitectate (rarely tectate, fossulate or intectate), with columellate infratectum, usually reticulate (sometimes spinulate or absent, rarely perforate).

Gynoecium Pistil composed of three connate carpels. Ovary inferior, usually trilocular (rarely unilocular). Style single, simple, filiform, with stylar canal (rarely trilobate). Stigma capitate, punctate or slightly trifid to deeply trilobate, usually papillate (with unicellular or multicellular papillae), usually Dry (sometimes Wet) type. Pistillodium absent.

Ovules Placentation usually axile (rarely basal). Ovules approx. twelve to more than 50 per carpel, anatropous, usually bitegmic (in Amaryllideae unitegmic; in some species of Crinum probably ategmic), usually crassinucellar (in Zephyranthes tenuinucellar). Micropyle bistomal (sometimes Z-shaped, zig-zag). Outer integument usually two (sometimes three or more) cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell or absent. Parietal tissue absent. Hypostase usually absent. Periclinal cell divisions often occurring in megasporangial epidermis (parietal cell not formed). Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Synergids with a filiform apparatus. Endosperm development usually helobial (sometimes nuclear; in Crinum cellular). Endosperm haustoria? Embryogenesis asterad or onagrad. Polyembryony occurring in at least Crinum and Hymenocallis.

Fruit Usually a loculicidal capsule (in, e.g., Boophone and Cybistetes irregularly dehiscent; in Calostemmateae indehiscent; in, e.g., Gethyllideae a berry).

Seeds Aril absent. Seed coat testal, sometimes fleshy or winged, sometimes (in, e.g., Galanthus and Leucojum) with caruncular elaiosome at chalazal end. Exotesta usually with thin phytomelan layer on epidermal cell walls (absent in water-rich seeds of Amaryllideae, Calostemmateae, Haemantheae, and Hymenocallideae, having fleshy seed coat and starchy endosperm). Endotesta? Tegmen collapsed. Perisperm not developed. Endosperm copious, with oils, aleuronic, starch or hemicellulose (sometimes rich in water), sometimes thin-walled, in Amaryllideae chlorophyllous. Embryo straight, small, little differentiated, with or without chlorophyll. Cotyledon one, bifacial, usually photosynthesizing. Cotyledon hyperphyll elongate, dorsiventrally flattened and assimilating, or compact and not assimilating. Hypocotyl internode absent. Coleoptile absent. Radicula well developed, contractile. Germination phanerocotylar.

Cytology n = 5–13, 20–23 or more (n = 52–55 in Ismene narcissiflora, n = 69 in Eucharis caucana, n = 59–89 in Sprekelia formosissima); x = 11 – Polyploidy frequently occurring. Chromosomes (1,5–)3–28 µm long. Agamospermy occurring in Crinum, Habranthus and Zephyranthes.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols (kaempferol, quercetin), norbelladine alkaloids (toxic tyrosine derivatives, more than 200 structures known at present, at least 79 of which from Narcissus; e.g. crinine, belladine, narwedine, galanthamine, haemanthamine, homolyrine, lycorenine, lycorine, homolycorine, pancratiostatine, and tazettine), benzylisoquinoline alkaloids, cyanogenic compounds, chelidonic acid and non-protein amino acids present. Mannans present as storage carbohydrates. Ellagic acid, proanthocyanidins, steroidal saponins, and allyl disulfides not found. Lectins binding mannose.

Use Ornamental plants, medicinal plants.

Systematics Amaryllidaceae are sister-group to Alliaceae.

Amaryllideae are sister to the remaining Amaryllidaceae, according to Meerow & al. (1999).

The variation in sieve tube plastids is large.

Amaryllideae Dumort., Anal. Fam. Plant.: 58. 1829

10–11/c 160. Amaryllis (1–2; A. belladonna; Northern and Western Cape), Nerine (23; southern Africa), Brunsvigia (20; southern Africa), Crossyne (2; Northern and Western Cape), Hessea (13; Namibia, Northern and Western Cape; incl. Namaquanula?), Namaquanula (2; southwestern Namibia, northernmost Northern Cape; in Hessea?), Strumaria (24; southern Africa, especially Namibia and southwesternmost South Africa), Boophone (6; tropical and southern Africa), Crinum (c 65; pantropical, southwards to South Africa), Ammocharis (5; tropical and southern Africa), Cybistetes (1; C. longifolia; Namibia, Northern and Western Cape). – Pantropical, with their highest diversity in South Africa. Bulb scales forming strongly elastic cotton-like fibres, when injured (due to presence of foliar fibres). Leaves with extensive fibres (with spiral cell wall thickenings); leaves developed prior to flowers (leaves in many Crininae perennial). Stomata paracytic; subsidiary cells with oblique divisions. Filaments usually connate at base (in Crininae free), sometimes with small processes (filament bases in Strumaria and Carpolyza adnate to style). Pollen grains usually disulcate. Exine intectate, columellate, often gemmate, with scattered spinulae. Style sometimes laterally inserted. Ovules unitegmic (sometimes ategmic?). Megagametophyte Allium type. Seeds watery, without resting period. Testa up to 25 cell layers thick, usually with chlorophyll and anomocytic stomata (absent in Amaryllis) or more or less collapsing, without phytomelan, or absent (e.g. in Crinum). Endosperm with starch and sometimes (e.g. in Crinum) with a suberine chlorophyllous layer. Embryo with chlorophyll. Very long tubular cotyledon envelope sometimes formed during germination. n = 10, 12, 15. Chromosomes 5,3–20,5 µm long. – Amaryllis and Boophone are successive sister-groups to the remaining Amaryllideae (Meerow & al. 1999).

[Cyrtantheae+[[Calostemmateae+Haemantheae]+[[Lycorideae+[Pancratieae+Narcisseae]]+ [Griffinia+ Worsleya+[Hippeastreae+[Eustephieae+[Hymenocallideae]]]]]]]

Vascular bundles with parenchymatic envelope cells.

Cyrtantheae Traub in Herbertia 5: 111. Nov 1938

1/c 50. Cyrtanthus (c 50; tropical and southern Africa, especially South Africa). – Peduncle without sclerenchyma cylinder. Collenchyma subepidermal. Rhizodermis one-layered. Velamen absent. Seeds flat, horizontally packed. Phytomelan present. n = (7) 8 (11).

[[Calostemmateae+Gethyllideae]+[[Lycorideae+[Pancratieae+Narcisseae]]+[Griffinia+ Worsleya+[Hippeastreae+[Eustephieae+[Hymenocallideae]]]]]]

[Calostemmateae+Gethyllideae]

Fruit indehiscent.

Calostemmateae D. Müll.-Doblies et U. Müll.-Doblies in Feddes Repert. 107: 7. Dec 1996

2/5–6. Calostemma (2; eastern Australia), Proiphys (3–4; Malesia, tropical and eastern Australia). – Malesia, tropical and eastern Australia. Bulbils as dispersal units. Ovules two or three per carpel. Embryo slowly germinating and producing a bulbil. Phytomelan absent. n = 10.

Gethyllideae Dumort., Anal. Fam. Plant.: 58. 1829

6/c 80. Clivia (5; South Africa, Swaziland), Cryptostephanus (5; tropical Africa), Scadoxus (9; tropical and southern Africa), Haemanthus (22; southern Africa, with their highest diversity in Namaqualand and Western Cape), Apodolirion (6; South Africa, Swaziland), Gethyllis (32; Namibia, Northern, Western and Eastern Cape, Free State, North-West). – Tropical and southern Africa (with their largest diversity in South Africa). Sometimes with rhizome. Rhizoderm one-layered. Velamen absent. Peduncle without sclerenchyma cylinder. Subepidermal collenchyma present. Inflorescence bracts usually connate. Gethyllis has up to 18 stamens. Fruit a berry. Seeds angular. Phytomelan usually absent. n = 6, 8, 9, 11, 12, 14. Alkaloids absent in Gethyllis.

[[Lycorideae+[Pancratieae+Narcisseae]]+[Griffinia+Worsleya+[Hippeastreae+[Eustephieae+ [Hymenocallideae]]]]]

[Lycorideae+[Pancratieae+Narcisseae]]

Seeds almost spherical, turgid.

Lycorideae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 235. 20 Jul 1943

2/c 26. Lycoris (c 30; China, the Korean Peninsula, Japan, Burma), Ungernia (6; Central Asia to Japan). – Temperate to subtropical East Asia to Iran, Central Asia and Burma. Seeds in Ungernia irregularly discoid. n = 11 or higher.

[Pancratieae+Narcisseae]

Pancratieae Dumort., Anal. Fam. Plant.: 58. 1829

2/c 30. Pancratium (20–21, the Canary Islands, the Mediterranean, West Africa, Namibia, southern Asia), Sternbergia (7–8; southeastern Europe, southwestern Asia, Kashmir). – Southeastern Europe, the Canary Islands, the Mediterranean, West Africa, Namibia, southwestern and southern Asia. Staminal tube dentate. n = 11.

Narcisseae Lam. et DC., Syn. Plant. Fl. Gall.: 165. 30 Jun 1806 [’Narcissi’]

7/65–75. Narcissus (30–40; Europe, the Mediterranean), Lapiedra (1; L. martinezii; Iberian Peninsula, western Mediterranean), Acis (9; the Mediterranean), Galanthus (19; Europe and eastwards to Iran), Leucojum (2; Europe and eastwards to Iran), Hannonia (1; H. hesperidium; Morocco), Vagaria (1–2; V. parviflora; Israel, Syria). – Europe, the Mediterranean, northwestern Africa, Crimea, Caucasus, western Asia to Iran. Peduncular bracts connate at base or along one side. Corona usually present in Narcissus. Elaiosome usually present. n = 7–9, 11, 12 (13) or higher. – Especially Galanthus have the inner tepal whorl very different from the outer one.

[Griffinia+Worsleya+[Hippeastreae+[Eustephieae+[Hymenocallideae]]]]

Rhizoderm one-layered. Velamen absent. Peduncle without sclerenchyma cylinder. Subepidermal collenchyma present. Bracts obvolute. Phytomelan frequently present.

Hippeastreae Herb. ex Sweet, Brit. Fl. Gard., ser. 2, 1: ad t. 14. 1 Sep 1829 [’Hippeastriformes’]

c 10/215–225. Griffinia (15–20; Brazil), Cearanthes (1; C. fuscoviolaceae; northeastern Brazil), Tocantinia (1; T. mira; Paraná in Brazil), 'Rhodophiala' (c 30; Brazil, Uruguay, Bolivia, Chile, Argentina; polyphyletic), Hippeastrum (c 55; Mexico, Central America, the West Indies, South America southwards to Bolivia and Argentina; incl. Worsleya?), Worsleya (1; W. procera; Organ Mountains in Brazil; in Hippeastrum?), ’Zephyranthes’ (c 70; southeastern United States, Central America, the West Indies, South America to Argentina; paraphyletic; incl. Habranthus and Rhodophila pro parte?), 'Habranthus' (35–40; Chile, Argentina; polyphyletic; in Zephyranthes?), Sprekelia (1–2; S. formosissima; Mexico, Guatemala), Traubia (1–2; T. modesta; Chile; incl. 'Rhodolirium' laetum?), Rhodolirium (2; Chile), 'Phycella' (c 8–10; Venezuela, Chile, Argentina; paraphyletic; incl. Placea?), 'Placea' (6; Chile; polyphyletic; in Phycella?). – Southeastern United States, Mexico, Central America, the West Indies, South America. Peduncular bracts often connate at base (sometimes only along one side). Flowers zygomorphic. Corona sometimes present. Stamens declinate, of various lengths. Seeds flat, winged or D-shaped. n = 8–13, 17 or more. Chromosomes 3–16.7 µm long. – Griffinia and Worsleya were each identified by Meerow & al. (1999) as an isolated clade in a basal trichotomy outside of Hippeastreae, Eustephieae and Hymenocallideae.

[Eustephieae+Hymenocallideae]

Mesophyll without palisade. Flowers actinomorphic. Tetraploidy often occurring.

Eustephieae Hutch., Fam. Fl. Plants 2: 130. 20 Jul 1934

3/14–16. Chlidanthus (6; southern Peru, Bolivia, northwestern Argentina), Eustephia (4; southern Peru, Bolivia), Hieronymiella (4–6; Bolivia, Argentina). – The Central Andes in Peru, Bolivia and Argentina. Stamens of two different lengths (dimorphic). Seeds flattened, winged.

Hymenocallideae Small, Man. S.E. Fl.: 315. 30 Nov 1933 [’Hymenocalleae’]

12–13/145–150. Hymenocallis (c 60; southeastern United States, the West Indies, northeastern South America), Leptochiton (2–3; the Andes), Stenomesson (c 13; the Andes), Clinanthus (22; the Andes in Ecuador to northwestern Argentina), Paramongaia (1; P. weberbaueri; Peru), Pamianthe (2; northern Andes), Phaedranassa (9; the Andes), Rauhia (2–3; northern Peru), Eucrosia (7; the Andes), Eucharis (17; tropical America; incl. Caliphruria?), Caliphruria (4; western Colombia, Peru; in Eucharis?), Plagiolirion (1; P. horsmannii; Cauca Valley in Colombia), Urceolina (5–7; southern central Peruvian Andes). – Southeastern United States, the West Indies, Central America, the Andes, tropical South America. Velamen present in Pamianthe. Leaves often petiolate, lorate. Flowers usually actinomorphic (sometimes zygomorphic). Androecial cup usually present. Pollen grains auriculate (the two ends tapering and with different sculpturing). Seeds flattened and obliquely winged, or spherical, turgid, and with glossy seed coat. Testa thick, spongy, vascularized, with chlorophyll. Phytomelan usually absent (present in Leptochiton). Embryo starchy. n = sometimes 19, 20, 22. Chromosomes 2.3–11.8 µm long.

Cladogram of Amaryllideae based on morphology and DNA sequence data (Meerow & Snijman 2001).

Cladogram (one of 5.000 successively weighted trees) of Amaryllidaceae based on DNA sequence data (Meerow & al. 1999).

ANEMARRHENACEAE Conran, M. W. Chase et Rudall

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Conran, Chase et Rudall in Kew Bull. 52: 995. Dec 1997

Genera/species 1/1

Distribution Northern China, the Korean Peninsula.

Fossils Unknown.

Habit Bisexual, perennial herb. Rhizome short, thick. Roots thick.

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vessels present in roots? Vessel elements with ? perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Idioblasts with calciumoxalate raphides sometimes present in foliar mesophyll.

Trichomes Hairs absent?

Leaves Alternate (spiral), simple, entire, linear, with ? ptyxis. Stipules absent; leaf sheath relatively well developed. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids? Leaf surface papillate. Leaf margin entire, papillate.

Inflorescence Terminal, spike-like panicle, with few-flowered partial inflorescences.

Flowers Actinomorphic, small. Pedicel articulated. Pericladium present. Hypogyny. Tepals 3+3, petaloid, more or less clawed, marcescent, usually free (sometimes partially connate at base). Septal nectaries with openings at stylar base. Disc absent.

Androecium Stamens three, antepetalous. Filaments flattened, free, adnate at middle of inner tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum probably secretory (pre-Ubisch-bodies present, resulting in Pollenkitt), with multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style short, subulate. Stigma small, capitate, type? Pistillodium absent.

Ovules Placentation axile. Ovules two per carpel, anatropous, apotropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument four to six cell layers thick. Inner integument two or three cell layers thick. Parietal cell formed from archesporial cell. Nucellar cap absent. Hypostase present. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent. Megagametophyte haustoria present. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule.

Seeds Aril absent. Seed coat exotestal. Exotesta with phytomelan layer on epidermal cell walls. Mesotesta, endotesta and tegmen collapsed? Perisperm not developed. Endosperm copious, fleshy, with pitted cell walls, with starch (with oil, aleurone, and hemicellulose?). Embryo relatively large, curved, chlorophyll? Cotyledon one (occasionally two?), well developed; cotyledon apex a haustorium hidden beyond seed coat. Cotyledon hyperphyll elongate. Hypocotyl internode absent. Coleoptile absent. Germination?

Cytology n = 11

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Steroidal saponins (sarsasaponigene, markogenin diglycoside, timosaponins A-III and B-II and Zhi-mu-, hinokiresinol-, smilageninoside- and anemarsaponin A1, A2 and B), glucanes (anemarin A, B, C and D), xanthone-C-glycoside and mangiferin present in rhizome.

Use Ornamental plant, medicinal plant.

Systematics Anemarrhena (1; A. asphodeloides; northern China, the Korean Peninsula).

Anemarrhena is sister to [Agavaceae+[Anthericaceae+[Behniaceae+Herreriaceae]]].

ANTHERICACEAE J. Agardh

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Agardh, Theoria Syst. Plant.: 27. Apr-Sep 1858 [’Anthericeae’]

Genera/species c 8/c 300

Distribution Europe, northern and eastern Africa, Madagascar, South and East Asia, northern and eastern Australia, southwestern North America to South America.

Fossils Unknown.

Habit Bisexual, perennial herbs (stem in Chlorophytum suffruticosum more or less lignified).

Vegetative anatomy Roots fibrous or thick and fleshy (sometimes with root nodules); velamen present at least in Anthericum and Chlorophytum; root exodermis sometimes thickened. Phellogen absent. Secondary lateral growth absent. Vessels present at least in roots (sometimes in stem). Vessel elements usually with simple (in stem also scalariform) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Styloids present in Chlorophytum. Mucilage cells and mucilage canals present at least in Echeandia. Tanniniferous cells absent. Silica bodies and laticifers absent. Calciumoxalate raphides usually abundant.

Trichomes Hairs?, usually absent.

Leaves Alternate (spiral or distichous), simple, entire, usually linear, with ? ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids as platelets, parallel to stomata (Convallaria type). Mesophyll with calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, thyrsoid, panicle or raceme-like (rarely umbel-like), with bostrychoid partial inflorescences, or raceme.

Flowers Usually actinomorphic (rarely zygomorphic). Pedicel usually articulated. Pericladium usually present. Hypogyny. Tepals 3+3, petaloid, often marcescent, usually free (in Diora and Leucocrinum connate at base into a tube). Septal nectaries present. Disc absent.

Androecium Stamens 3+3. Filaments usually free (in Echeandia connate), usually adnate at base (in Leucocrinum for most of their length) to tepals. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, usually with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular (in Chlorophytum stipitate, on gynophore). Style single, simple, filiform. Stigma capitate or trilobate, usually papillate (with unicellular papillae), Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules two to numerous per carpel, anatropous or campylotropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument approx. four cell layers thick. Inner integument two cell layers thick. Hypostase present. Parietal cell formed from archesporial cell. Nucellar cap in Leucocrinum formed by periclinal cell divisions in apical part of megasporangial epidermis. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus (Chlorophytum). Antipodal cells often persistent. Megagametophyte haustoria frequently present. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Frui A loculicidal capsule (rarely a schizocarp or a nut) often with persistent tepals.

Seeds Aril absent. Seed coat testal. Exotesta with phytomelan layer on epidermal cell walls. Outer testal layers sometimes with thickened periclinal cell walls. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, fleshy, with lipids and starch; endosperm cell walls with pits. Embryo cylindrical, incurved, without chlorophyll. Cotyledon one, short, with apex transformed into haustorium, not photosynthesizing, in Chlorophytum with a tubular elongation of cotyledon sheath. Cotyledon hyperphyll? Hypocotyl internode? Mesocotyl absent. Coleoptile present in Chlorophytum. Germination cryptocotylar.

Cytology n = 7, 8, 10, 11, 13–15 or more – Polyploidy frequently occurring. Chromosomes 2–10 (to 13.8) µm long.

DNA The mitochondrial gene rpl2 is absent (lost). A genome duplication has taken place in Chlorophytum.

Phytochemistry Steroidal saponins, phenylalanine-derived cyanogenic glycosides (in Chlorophytum), and chelidonic acid present. Flavonols, ellagic acid, alkaloids, and anthraquinones not found.

Use Ornamental plants.

Systematics Echeandia (75–80; southwestern United States, Mexico, Central America, northern South America to Peru), Anthericum (c 65; Europe, North and East Africa southwards to Tanzania, southwestern Asia), Paradisea (2; Iberian Peninsula, mountains in southern Europe), ‘Chlorophytum’ (c 150; Africa, Madagascar, India and eastwards to northern and eastern Australia; polyphyletic), Leucocrinum (1; L. montanum; southwestern United States); unplaced Diamena (1; D. stenantha; upper Cerro de las Cabras near Trujillo in Peru, possibly extinct), Diora (1; D. cajamarcaensis; the Andes in Peru), Eremocrinum (1; E. albomarginatum; Utah, northern Arizona), Hagenbachia (5; Central and South America).

Anthericaceae are sister-group to [Behnia+[Herreriaceae]].

The Echeandia clade is sister to the remaining Anthericaceae (Kim & al. 2010).

Cladogram (simplified) of Anthericaceae based on DNA sequence data (Kim & al. 2010).

APHYLLANTHACEAE Burnett

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Burnett, Outl. Bot.: 421, 1153. Feb 1835

Genera/species 1/1

Distribution Western Mediterranean from southern France to northern Morocco.

Fossils Unknown.

Habit Bisexual, perennial herbs. Xeromorphic, with photosynthesizing stems. Tufts consisting of long-scapose inflorescences.

Vegetative anatomy Roots fibrous; mature roots without cortex; all root tissues except phloem lignified. Rhizome stele differentiated into inner, primary, irregularly oriented traces and outer, secondary, serial vascular traces. Phellogen absent. Secondary lateral growth (in rhizome) anomalous. Vessels present in roots. Vessel elements with simple or scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type (in roots), with cuneate protein crystals. Nodes? Mucilage cells abundant. Idioblasts with calciumoxalate raphides frequent in rhizome cortex. Cuticular wax crystalloids on stems as non-orientated platelets.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, scale-like, reduced to non-photosynthesizing leaf sheath; uppermost leaf with rudimentary pseudolamina and with short ligule, with supervolute subinvolute ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous. Stomata absent from leaves. Mesophyll often with calciumoxalate raphides. Epidermal cells of abaxial side elongate, with thickened and lignified walls. Leaf margin entire.

Inflorescence Terminal, unbranched, one- or few-flowered capitate. Peduncle (scape) long and narrow, terete, photosynthesizing, with thickened epidermis and rows of deeply sunken anomocytic stomata in upper part; medullary parenchyma often collapsing and replaced by an air canal; cuticle thick; cuticular wax crystalloids as parallel rodlets (Convallaria type); calciumoxalate as raphides. Bracts dry, scale-like, membranous, one or two free and five connate.

Flowers Actinomorphic. Pedicel not articulated, very short, or absent. Hypogyny. Tepals 3+3, petaloid, clawed, marcescent, free. Septal nectaries infralocular. Disc absent.

Androecium Stamens 3+3. Filaments free from each other, adnate to base of tepals (epitepalous). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains spiraperturate, shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, echinate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple, with stylar canal. Stigma trilobate with short filiform lobes, papillate, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovule one per carpel, anatropous or somewhat campylotropous-amphitropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument three or four cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell. Megagametophyte monosporous, Polygonum type. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule with persistent style.

Seeds Aril absent. Seed coat exotestal. Exotesta with thin phytomelan layer on epidermal cell walls. Mesotesta, endotesta and tegmen more or less collapsed. Perisperm not developed. Endosperm copious, with lipids and aleurone (starch absent). Embryo large, straight, well differentiated, with chlorophyll. Cotyledon one, well developed; cotyledon leaf sheath with ligule. Cotyledon hyperphyll elongate, terete, assimilating. Hypocotyl internode short. Coleoptile absent. Germination cryptocotylar?

Cytology n = 16

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols, steroidal saponins (genin, 25D- and 25L-saponogene), and wax alcohols present. Chelidonic acid? Ellagic acid, proanthocyanidins, and alkaloids not found.

Use Unknown.

Systematics Aphyllanthes (1; A. monspeliensis; southern France, the Iberian Peninsula, northern Morocco).

In the analysis by Wurdack & Dorr (2009), Aphyllanthes is recovered as part of a trichotomy in the same clade as Hyacinthaceae, Themidaceae, Anemarhenaceae and the Agavaceae group.

ASPARAGACEAE Juss.

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de Jussieu, Gen. Plant.: 40. 4 Aug 1789 [’Asparagi’], nom. cons.

Asparagales Link, Handbuch 1: 272. 4-11 Jul 1829 [’Asparaginae’]; Asparagineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.]7: 1556. 1846 [‘Asparageae’]; Hemiphylacaceae Doweld, New Syllabus Plant Fam.: 921. Apr 2007

Genera/species 2/165–195(–c 300?)

Distribution Eurasia, Africa, Australia, Mexico.

Fossils Unknown.

Habit Usually bisexual (sometimes monoecious, andromonoecious, dioecious, or gynodioecious), evergreen or deciduous shrubs or suffrutices, or perennial herbs, often twining and climbing. Tuberous stem present in Asparagus ovatus and A. undulatus. Branches usually green and assimilating. Branch ends in Asparagus transformed into foliaceous or terete phyllocladia. Spines consisting of transformed branches or leaf bases present in some species.

Vegetative anatomy Lateral roots sometimes tuberous; sometimes with membranous multiple velamen; roots in Hemiphylacus contractile. Phellogen? Secondary lateral growth absent. Vessels present in roots and stem. Vessel elements with scalariform (in roots often also simple) perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays?, Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Idioblasts (raphide cells) with calciumoxalate raphides.

Trichomes Hairs usually absent (rarely unicellular?).

Leaves Alternate (spiral), simple, entire, needle- or scale-like (in Hemiphylacus linear), with basal spur (sometimes transformed into spine), with ? ptyxis. Stipules and leaf sheath absent. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids as parallel platelets (Convallaria type) or absent. Mesophyll usually with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin entire. Extrafloral nectaries present in some species.

Inflorescence Axillary, fasciculate, panicle or umbel-like, or flowers solitary (in Hemiphylacus raceme or thyrse). Floral prophylls (bracteoles?) in Hemiphylacus lateral.

Flowers Actinomorphic, usually small. Pedicel articulated; pericladium separated from pedicel by a swelling. Hypogyny. Tepals 3+3, petaloid, free or connate in lower part or at base into a tube (in Hemiphylacus connate half-way up, twisted following anthesis). Septal nectaries present. Disc absent.

Androecium Stamens usually 3+3 (in Hemiphylacus three fertile stamens and three staminodia). Filaments filiform or flattened, free from each other, adnate at base to tepals; in Hemiphylacus three inner antepetalous fertile stamens adnate to inner tepals, and three outer staminodia adnate to outer tepals. Anthers usually basifixed (in Hemiphylacus dorsifixed), versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers with staminodia. In Hemiphylacus flowers bisexual, with three staminodia.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, starchy, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate? infratectum, psilate-microperforate or somewhat reticulate.

Gynoecium Pistil composed of three connate carpels; carpels in Hemiphylacus antepetalous. Ovary superior, trilocular (sometimes slightly stipitate). Style single, simple, with stylar canal. Stigma capitate to trilobate, Dry or Wet type. Male flowers with pistillodium.

Ovules Placentation axile. Ovules two to twelve (in Hemiphylacus three to six) per carpel, hemianatropous or almost orthotropous (in Hemiphylacus campylotropous), bitegmic, crassinucellar. Micropyle at least in Hemiphylacus endostomal. Outer integument five to seven cell layers thick. Inner integument two cell layers thick. Obturator present near micropyle in Hemiphylacus. Hypostase present in Hemiphylacus. Parietal cell in Hemiphylacus formed from archesporial cell. Micropylar megasporangial epidermal cells enlarged and forming a uniseriate layer, crushed in mature seed (also in Hemiphylacus). Nucellar cap absent. Megagametophyte monosporous, Polygonum type. Megagametophyte curved and asymmetrical (also in Hemiphylacus). Endosperm development usually nuclear (in Hemiphylacus helobial). Endosperm haustoria? Embryogenesis?

Fruit Usually a one- or several-seeded berry (rarely a nut; in Hemiphylacus a loculicidal capsule).

Seeds Aril absent. Pseudo-operculum sometimes present. Seed coat exotestal. Testa multiplicative. Exotesta massive, with phytomelan layer on epidermal cell walls (always?). Mesotesta and endotesta collapsed. Tegmen inconspicuous, collapsed. Perisperm not developed. Endosperm copious, with oil, aleurone and hemicellulose (starch absent); endosperm cells thick, with pitted walls. Embryo large, straight to curved, well differentiated, without chlorophyll. Cotyledon one. Cotyledon hyperphyll? Hypocotyl internode short. Coleoptile absent. Radicula persistent. Germination?

Cytology n = 10 (Asparagus), n = 56 (Hemiphylacus) – Polyploidy frequently occurring in Asparagus. Sex chromosomes present in, e.g., Asparagus officinalis. Chromosomes in Hemiphylacus 1–3 µm long.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols (kaempferol, quercetin), steroidal saponins, acetidine carbonic acid, and chelidonic acid present. Carbohydrates stored as inulin-like fructans and mannans. Ellagic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found.

Use Ornamental plants, vegetables (Asparagus officinalis, A. albus etc.).

Systematics Asparagus (160–190; mainly arid and subarid regions in Europe, Africa, Madagascar, the Mascarene Islands, Asia, and one species in Australia), Hemiphylacus (5; semiarid regions in central Mexico).

Asparagaceae are sister to Ruscaceae.

ASPHODELACEAE Juss.

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de Jussieu, Gen. Plant.: 51. 4 Aug 1789 [‘Asphodeli’], nom. cons.

Aloaceae Batsch, Tab. Affin. Regni Veg.: 138. 2 Mai 1802 [‘Alooideae’]; Asphodelales Doweld, Tent. Syst. Plant. Vasc.: lvii. 23 Dec 2001; Asphodelineae Thorne et Reveal in Bot. Rev. (Lancaster) 73: 80. 29 Jun 2007

Genera/species 18–19/760–785

Distribution Southern and eastern Europe, the Mediterranean, Africa, Madagascar, Socotra, the Arabian Peninsula, the Mascarene Islands, West and Central Asia, Australia, New Zealand, with the largest diversity in South Africa.

Fossils Uncertain.

Habit Bisexual, usually perennial herbs (some species of Aloe and Kniphofia are pachycaul shrubs or trees). Rarely with bulb. Many representatives are leaf succulents. Roots often somewhat succulent, sometimes contractile, sometimes swollen.

Vegetative anatomy Root velamen present in some species. Stem usually fibrous. Phellogen? Secondary lateral growth anomalous (present especially in Aloe, but also in Bulbine, Kniphofia and Trachyandra) or absent. Vessels present at least in roots (in some species also in stem). Vessel elements in roots usually with simple perforation plates, in stem with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals, or (in Alooideae) P2cf type, with cuneate protein crystals and peripheral protein filaments. Nodes? Mucilage cells with calciumoxalate as raphides or prismatic crystals.

Trichomes Hairs absent?

Leaves Alternate (spiral or distichous), simple, entire, often linear or subulate, in many species succulent (some species have ‘window leaves’ with transparent tissue in distal part), with ? ptyxis. Stipules absent; leaf sheath usually closed (sometimes absent). Venation parallelodromous; veins often indistinct; vascular bundles often inverted; large group of thin-walled parenchymatous (secretory?) aloin cells present adjacent to phloem in envelope of inner vascular bundle (in, e.g., Kniphofia these cells with lignified walls). Stomata usually anomocytic (rarely paracytic; in Alooideae tetracytic). Cuticular wax crystalloids as non-orientated rodlets or platelets. Mesophyll with mucilaginous idioblasts containing calciumoxalate as raphides or single prismatic crystals. Leaf margin serrate or entire. Leaf apex often pointed.

Inflorescence Terminal, panicle, raceme- or spike-like, often branched.

Flowers Usually actinomorphic (in Haworthia and few species of Aloe weakly zygomorphic, bilabiate, in Gasteria curved). Pedicel in Asphodelus, Asphodeline and Bulbine articulated. Hypogyny. Tepals 3+3, petaloid, usually free (in Kniphofia and Aloeae connate into a tubular or campanulate perianth). Septal nectaries well developed, infralocular. Disc absent.

Androecium Stamens 3+3. Filaments subulate to filiform, free from each other and from tepals (in Bulbine with numerous long hairs). Anthers usually dorsifixed (in Eremurus basifixed; filaments often? inserted in a connective pit), versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory (tapetal material slightly invading anther locule at tetrad stage in Aloe). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate or trichotomosulcate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate or reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple. Stigma punctate, usually Dry (sometimes Wet) type. Pistillodium absent.

Ovules Placentation axile. Ovules one to more than 40 per carpel, anatropous, orthotropous (Asphodeleae), hemitropous or campylotropous, bitegmic, crassinucellar. Micropyle usually endostomal? (in Kniphofia bistomal). Outer integument three or four cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell, forming parietal tissue between megasporocyte and megasporangial epidermis. Hypostase present. Megasporangial endothelium present in Kniphofia. Megagametophyte monosporous, Polygonum type. Synergids in Asphodelus with a filiform apparatus. Endosperm development ab initio helobial. Endosperm haustoria? Embryogenesis onagrad, asterad or chenopodiad.

Fruit A loculicidal capsule (in some species of Aloe a fleshy berry-like capsule).

Seeds Seeds sometimes winged. Arilloid structure formed as annular invagination at distal part of funicle (seemingly from an extra ‘integument’), two or several cell layers thick. Seed coat exotestal. Exotesta usually with phytomelan layer on epidermal cell walls. Endotesta and tegmen collapsed. Perisperm usually absent (sometimes sparsely developed). Endosperm copious, thick-walled, with lipids and aleuronic (sometimes with hemicellulose?). Embryo long, straight, well differentiated, without chlorophyll. Cotyledon one. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode present or absent. Mesocotyl absent. Coleoptile sometimes present. Germination cryptocotylar?

Cytology n = 2–7, 12–16, 20, 40, 44, 49; x = 7 (Aloeae) – Karyotype in Aloeae bimodal, in Bulbine etc. with four long and three short chromosomes. Chromosomes 1,5–20 µm long.

DNA The mitochondrial gene rpl2 is absent. Loss of the 3’-rps12-intron (Bulbine?).

Phytochemistry Flavonoids, alkaloids, cyanogenic substances, anthraquinones (1-methyl-8-hydroxyanthraquinones in roots), anthrones (e.g. anthrone-C-glycoside [aloin, barbaloin] in leaves), tetrahydroanthracenones (chrysophanol, 10,7’-bichrysophanol, knipholone, asphodeline and other lipophilic anthranoid aglucones), chromones, phenylpyrones, polyacetate derived arthroquinones, phenolic amines, chelidonic acid, bitter substances, and resins present. Mannans present as storage carbohydrates. Flavonols, ellagic acid, proanthocyanidins, and steroidal saponins not found.

Use Ornamental plants, medicinal plants and cosmetics (Aloe vera, A. ferox etc.).

Systematics Asphodelaceae are probably sister-group to [Hemerocallidaceae+Xanthorrhoeaceae] (or, alternatively, to Xanthorrhoea).

A probable topology is [Asphodeleae+[Kniphofieae+Aloeae]] (Treutlein & al. 2003).

Asphodeleae Lam. et DC., Syn. Plant. Fl. Gall.: 160. 30 Jun 1806 [‘Asphodeli’]

2/26. Asphodelus (12; the Mediterranean, southwestern Asia), Asphodeline (16; the Mediterranean and eastwards to Caucasus). – The Mediterranen, southwestern Asia. – Asphodeleae are sister to all other Asphodelaceae.

[Kniphofieae+Aloeae]

Kniphofieae Hutch., Fam. Fl. Plants 2: 83, 90. 20 Jul 1934

4/c 180. Eremurus (c 45; eastern Europe, alpine regions in West and Central Asia), Trachyandra (c 50; tropical and southern Africa, Madagascar, with their highest diversity in Western Cape); Bulbinella (22; Northern, Western and Eastern Cape, New Zealand), Kniphofia (65–70; tropical and southern Africa, Madagascar, the Arabian Peninsula). – Eastern Europe to Central Asia, tropical and southern Africa, Madagascar, the Arabian Peninsula, New Zealand. Aloin cells are absent and replace by a well developed sclerenchymatic cap (n.b. also in some Aloeae). Aloin cells have also been reported from Dianella (Hemerocallidaceae). Bulbinella and Kniphofia have knipholone.

Aloeae A. Rich. in C. H. D. d’Orbigny, Dict. Univ. Hist. Nat. 1: 291. 18 Dec 1841 [‘Aloineae’]

12–13/560–585. Bulbine (50–60; tropical and southern Africa, Australia, with their highest diversity in Western Cape; incl. Jodrellia?), Jodrellia (3; Central to northeastern Africa; in Bulbine?); Aloidendron (6–7; southern Africa, Somalia), Kumara (2; Western Cape), Haworthia (c 40; Namibia, South Africa, with their highest diversity in Western and Eastern Cape), Aloiampelos (7; southern and eastern South Africa, especially Western and Eastern Cape, Swaziland), Aloe (400–420; tropical, eastern and southern Africa, Madagascar, the Arabian Peninsula, Socotra), Gasteria (25–30; Namibia, South Africa, Mozambique), Haworthiopsis (18; Western Cape), Astroloba (6; Western and Eastern Cape), Aristaloe (1; A. aristata; mountains in southeastern South Africa), Gonialoe (3; Namibia, arid regions in South Africa), Tulista (4; Western Cape). – Africa, Madagascar, the Arabian Peninsula, Socotra, Australia, with their largest diversity in South Africa. Sometimes shrub-like. Sieve tube plastids P2cf type, with cuneate protein crystals and peripheral protein filaments. Stomata usually tetracytic. Foliar vascular bundles forming a cylinder; globuli present in outer vascular bundle envelope (also present in Kniphofia). Karyotype bimodal. 1-methyl-8-hydroxy anthraquinones (e.g. chrysophanol) present in roots and anthrone-C-glycosides in leaves. – Bulbine (including Jodrellia?) is sister to the remaining Aloeae. Bulbine has knipholone and a bimodal karyotype based on n = 7: four long and three short chromosomes.

Phylogeny (simplified) of Asphodelaceae based on DNA sequence data (Treutlein & al. 2003; Daru & al. 2012; Grace & al. 2013).

Phylogeny of Asphodelaceae based on Manning & al. (2014).

ASTELIACEAE Dumort.

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Dumortier, Anal. Fam. Plant.: 59, 61. 1829

Asteliales Dumort., Anal. Fam. Plant.: 59. 1829 [‘Astelarieae’]

Genera/species 3/36

Distribution The Mascarene Islands, New Guinea, New Caledonia, Fiji, Samoa, the Society Islands, the Marquesas Islands, Hawaiian Islands, temperate regions in southeastern Australia, Tasmania, New Zealand, Auckland, Campbell and Chatham Islands, southern Chile and Argentina, the Falkland Islands.

Fossils Pollen of ‘Astelia’ is known from the Late Eocene onwards in New Zealand.

Habit Bisexual (Milligania), dioecious or gynodioecious, perennial herbs. Rarely epiphytic.

Vegetative anatomy 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 P2c type, with cuneate protein crystals, or P2cs type, with cuneate protein crystals and starch grains. Nodes? Idioblasts with calciumoxalate raphides frequent. Crystals of various shape present in Astelia and Collospermum.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, stellate or lepidote, often long and with multicellular base, marginal cells often caducous.

Leaves Alternate (spiral), simple, entire, often linear, with ? ptyxis. Stipules absent; leaf sheath open or closed (sometimes absent). Venation parallelodromous; often with two distinct lateral veins (often larger than midvein) and, in connection with these, a well developed sclerenchyma. Stomata paracytic. Cuticular wax crystalloids as parallel platelets (Convallaria type). At least young leaves densely pubescent. Adaxial hypodermis present. Mesophyll with calciumoxalate as raphides and single prismatic crystals. Some species of ‘Astelia’ and Collospermum with secretory cavities (mucilaginous canals). Leaf margin entire.

Inflorescence Terminal, branched system of racemes or spikes. Bracts large. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic. Usually hypogyny (in Milligania half epigyny). Tepals usually 3+3 (in Neoastelia spectabilis sometimes 5+5 or 7+7), sepaloid, petaloid or bract-like, membranous or fleshy, usually connate at base (sometimes free). Septal nectaries branched, in furrows on ovary, sometimes external, on abaxial side of ovary (in Milligania short, in upper part of ovary). Disc absent.

Androecium Stamens 3+3. Filaments usually free (in Milligania adnate at base to tepals). Anthers basifixed to dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Female flowers with staminodia.

Pollen grains Microsporogenesis usually successive (in Milligania simultaneous). Pollen grains usually monosulcate (sometimes trichotomosulcate), shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, foveolate, echinate, spinulate, or smooth.

Gynoecium Pistil composed of three (to seven) connate carpels. Ovary usually superior (in Milligania semi-inferior), usually trilocular (in some species of ‘Astelia’ unilocular), in many species of ‘Astelia’ and Collospermum with mucilage hairs on inner side (from funicles and placentae). Stylodia usually absent (in Milligania short). Stigma usually trilobate, decurrent (in some species of ‘Astelia’ connate into a wide stigmatic surface), papillate, Dry type. Male flowers with pistillodium (sometimes with fertile ovules).

Ovules Placentation usually axile (in some species of ‘Astelia’ parietal). Ovules four to c. 15? per carpel, anatropous, bitegmic, pseudocrassinucellar. Funicle hairy in many species (not in Milligania). Micropyle bistomal, also Z-shaped (zig-zag). Outer integument two to five cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell, degenerating (parietal tissue absent). Hypostase present. Nucellar cap absent. Megagametophyte monosporous, Polygonum type, with chalazal constriction. Antipodal cells persistent. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit Usually a berry (in Milligania a loculicidal capsule dehiscing from apex).

Seeds Aril absent. Seeds in Collospermum surrounded by mucilage hairs. Seed coat exotestal. Testa hard. Exotesta with phytomelan layer on epidermal cell walls. Endotesta collapsed? Tegmen in ‘Astelia’ reduced to a layer of empty cells. Perisperm not developed. Endosperm copious, with oil and aleurone (starch and hemicellulose absent). Embryo straight or somewhat curved, well differentiated, chlorophyll? Cotyledon one. Cotyledon hyperphyll elongate to compact, often assimilating. Hypocotyl internode short. Coleoptile absent. Radicula well developed, ephemeral. Germination?

Cytology n = 8, 30, 35, 105 (x = 8) – Polyploidy occurring. Chromosomes 4–6 μm long.

DNA

Phytochemistry Flavonols (kaempferol, quercetin, isorhamnetin), steroidal saponins, and chlorogenic acid present. Ellagic acid and proanthocyanidins not found.

Use Unknown.

SystematicsAstelia’ (30; the Mascarene Islands, New Guinea, New Caledonia, temperate parts of southeastern Australia, Tasmania, New Zealand, Auckland, Campbell and Chatham Islands, Vanuatu, Fiji, Samoa, the Society Islands, the Marquesas Islands, the Hawaiian Islands, southern Chile and Argentina, the Falkland Islands), Neoastelia (1; N. spectabilis; northeastern New South Wales), Milligania (5; Tasmania).

Collospermum is nested within Astelia. Milligania and Neoastelia are sister-groups.

In a study by Kocyan & al. (2011), Asteliaceae form a paraphyletic grade basal to Blandfordia (Blandfordiaceae), Lanaria (Lanariaceae) and Hypoxidaceae.

Bayesian inference 50% consensus tree (simplified) of Asteliaceae based on plastid and nuclear DNA (Birch & al. 2012).

BEHNIACEAE Conran, M. W. Chase et Rudall

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Conran, Chase et Rudall in Kew Bull. 52: 996. Dec 1997

Genera/species 1/1

Distribution Southeastern Africa.

Fossils Unknown.

Habit Dioecious (cryptodioecious), perennial herb with more or less lignified stems, occasionally somewhat twining or climbing. Xeromorphic. Aerial roots absent.

Vegetative anatomy Roots fibrous; velamen one-layered; root exodermis uniseriate. Phellogen? Stem cortex with a cylinder of fibre bundles and small collateral vascular bundles; centre of stem with scattered bundles. Rhizome with lignified secondary xylem, parenchymatous secondary phloem and anomalous secondary lateral growth. Vessels present in roots and stem. Vessel elements with simple or scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Mucilage cells and laticifers absent. Tanniniferous cells absent. Silica bodies absent. Calciumoxalate raphides abundant; druses sometimes present.

Trichomes Hairs absent?

Leaves Alternate (distichous), simple, entire, linear, with supervolute? ptyxis, differentiated into pseudopetiole and pseudolamina. Stipules and leaf sheath absent. Venation pinnate-parallelodromous, acrodromous; midvein distinct, longitudinal main veins connected with parallel transverse secondary veins. Stomata paracytic; subsidiary cells with oblique division. Cuticular wax crystalloids? Mesophyll with calciumoxalate raphides. Leaf margin entire.

Inflorescence Axillary, bostryx, or flowers solitary.

Flowers Actinomorphic, small. Pedicel articulated. Pericladium short. Hypogyny. Tepals 3+3, petaloid, marcescent, connate into a tube. Septal nectaries present (also in male flowers). Disc absent.

Androecium Stamens 3+3. Filaments filiform, free from each other, adnate in lower part to tepals (epitepalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Female flowers with staminodia.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, spinulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular, on gynophore. Style single, simple. Stigma trilobate, papillate, Wet type. Male flowers with pistillodium.

Ovules Placentation axile. Ovules two or three per carpel, anatropous, bitegmic, tenuinucellar (crassinucellar?). Micropyle endostomal. Outer integument three or four cell layers thick. Inner integument two cell layers thick. Hypostase present. Parietal cell formed from archesporial cell. Parietal tissue absent. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A few- to many-seeded berry.

Seeds Aril absent. Testal and tegmic cells thin-walled. Phytomelan layer probably absent. Exotesta collapsed, sometimes caducous. Endotesta parenchymatous, partially collapsed. Tegmen crushed. Perisperm not developed. Endosperm copious, with oil and aleurone (starch and hemicellulose absent); endosperm cells walls not pitted. Embryo short, straight, chlorophyll? Cotyledon one, little differentiated (ligule absent). Cotyledon hyperphyll? Hypocotyl internode? Coleoptile absent? Germination?

Cytology n = ?

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Very insufficiently known. Steroidal saponins present.

Use Unknown.

Systematics Behnia (1; B. reticulata; Zimbabwe, South Africa, Swaziland).

Behnia is sister to Herreriaceae.

BLANDFORDIACEAE R. Dahlgren et Clifford

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Dahlgren et Clifford in Dahlgren, Clifford et Yeo, Fam. Monocot.: 155. 1985

Genera/species 1/4

Distribution Eastern Australia, Tasmania.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome tuberous.

Vegetative anatomy Roots fibrous, with uniseriate velamen. Root cortex starchy. Phellogen absent. Stem cortex collenchymatous, reduced, with a thick cylinder of lignified fibre bundles; inner cortex with small collateral bundles; centre of stem hollow. 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 P2c, with cuneate protein crystals. Nodes? Calciumoxalate druses present. Raphides and styloids absent.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, linear, with flat, curved or somewhat concave ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous; midvein prominent. Stomata anomocytic. Cuticular wax crystalloids? Secretory cavities absent. Mesophyll often with cells containing calciumoxalate druses (raphides absent). Leaf margin entire.

Inflorescence Usually terminal, simple raceme; flowers sometimes solitary axillary. Floral prophylls (bracteoles) two.

Flowers Actinomorphic, large. Pedicel articulated. Pericladium long, pedicel-like. Hypogyny. Tepals 3+3, petaloid, persistent, connate into a long tube. Septal nectaries external, as deep furrows on abaxial side of ovary and gynophore. Disc absent.

Androecium Stamens 3+3. Filaments filiform, free from each other, adnate in lower part to perianth tube (epitepalous). Anthers centrifixed (seemingly basifixed), versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits); connective hollow in lower part and sheathing filament. Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains trichotomosulcate, shed as monads, bicellular at dispersal. Exine tectate? with ? infratectum, perforate?, verrucate?

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular, stipitate (on gynophore). Style single, simple, short. Stigma entire, somewhat trilobate, punctate, with three furrows, papillate?, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules c. 40 to c. 50 per carpel, anatropous, bitegmic, pseudotenuinucellar (functionally tenuinucellar). Micropyle endostomal. Outer integument four or five cell layers thick, somewhat shorter than inner integument. Inner integument two cell layers thick. Parietal cell formed from archesporial cell, early degenerating (parietal tissue absent). Hypostase present. Nucellar cap approx. two cell layers thick, formed by periclinal divisions of epidermal cells of megasporangium. Megagametophyte monosporous, Polygonum type, with chalazal constriction. Synergids with a filiform apparatus. Antipodal cells persistent, sometimes becoming binucleate. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule with accrescent gynophore.

Seeds Aril absent. Seed coat exotestal. Exotesta with papillar hairs, without phytomelan. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, with oil and aleurone (without starch). Embryo short, somewhat curved, well differentiated, chlorophyll? Cotyledon one, as a photosynthesizing hyperphyll without leaf sheath. Hypocotyl internode? Coleoptile absent? Germination phanerocotylar.

Cytology n = 17, 27

DNA

Phytochemistry Flavonols (kaempferol, quercetin), flavone-C-glycosides, cyanidin, and chelidonic acid present. Steroidal saponins? Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Blandfordia (4; eastern Australia from southeastern Queensland to Tasmania).

A probable topology is [Blandfordia+[Asteliaceae+[Hypoxidaceae+Lanaria]]].

BORYACEAE (Baker) M. W. Chase, Rudall et Conran

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Chase, Rudall et Conran in Kew Bull. 52: 416. Jul 1997

Genera/species: 2/12

Distribution: Australia.

Fossils: Unknown.

Habit: Bisexual, perennial herbs. Xerophytes. Roots in Borya tuberculate and slightly lignified (with Rhizoctonia or other fungi?), sometimes stilt roots.

Vegetative anatomy: Roots fibrous, with mycorrhiza, Root hypodermis lignified; endodermis strongly thickened; inner cortex sclerenchymatous; medulla lignified. Phellogen absent. Stem endodermis in Borya strongly lignified; vascular strands in peripheral cylinder inside endodermis; medulla lignified, with fibres (in Alania parenchymatous). Secondary lateral growth usually absent (older stems in Borya with anomalous secondary lateral growth). 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 P2c type, with cuneate protein crystals. Nodes? Calciumoxalate as raphides.

Trichomes: Hairs absent?

Leaves: Alternate (spiral), simple, entire, linear to filiform, with ? ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous. Vascular bundles containing lateral phloem. Stomata anomocytic. Cuticular wax crystalloids? Hypodermis with lignified fibres. Inner epidermal cell walls lignified. Leaf margin finely serrate or entire.

Inflorescence: Terminal, raceme- or spike-like. Peduncle (scape) long. Inflorescence surrounded by an involucre; involucral bracts in Alania dry, in Borya differentiated into subulate foliaceous outer and scale-like inner bracts. Pedicel in Alania with several floral prophylls (bracteoles), in Borya with a single floral prophyll.

Flowers: Actinomorphic. Hypogyny. Tepals 3+3, petaloid, persistent, free (Alania), or connate in lower part into a short tube (Borya). Septal nectaries external-basal, as deep furrows in ovary wall. Disc absent.

Androecium: Stamens 3+3. Filaments filiform, free from each other, in Borya adnate to perianth tube (epitepalous). Anthers almost as wide as long, sometimes glandular, usually basifixed (sometimes centrifixed), non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

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

Gynoecium: Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple, filiform (in Alania geniculate). Stigma capitate, type? Pistillodium absent.

Ovules: Placentation axile. Ovules numerous per carpel, anatropous, bitegmic, pseudotenuinucellar (functionally tenuinucellar, developmentally crassinucellar). Micropyle bistomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell, early degenerating (parietal tissue absent). Hypostase present. Megasporangial tissue one cell layer thick. Nucellar cap absent. Megagametophyte monosporous, Polygonum type, with chalazal constriction. Synergids without filiform apparatus. Antipodal cells persistent. Endosperm development ab initio helobial. Endosperm haustoria absent. Embryogenesis?

Fruit: A loculicidal capsule with persistent perianth.

Seeds: Aril absent. Seeds in Borya papillate. Seed coat exotestal. Exotesta with phytomelan layer on epidermal cell walls. Exotestal cells with thickened walls. Endotesta? Tegmen degenerating. Hypostase somewhat lignified. Perisperm not developed. Endosperm copious, with oil and aleurone (starch absent). Embryo small, ovoid, chlorophyll? Cotyledon one. Cotyledon hyperphyll? Hypocotyl internode? Coleoptile? Germination?

Cytology: n = 11 (Alania); n = 14, 28 (Borya)

DNA:

Phytochemistry: Very insufficiently known. Chelidonic acid present.

Use: Unknown.

Systematics: Borya (11; Western Australia, Queensland, Victoria), Alania (1; A. endlicheri; mountains west of Sydney in New South Wales).

Boryaceae are sister to the clade [Blandfordia+[Asteliaceae+[Lanaria+Hypoxidaceae]]]. Sometimes, Boryaceae have been recovered as sister-group to Orchidaceae.

DORYANTHACEAE R. Dahlgren et Clifford

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Dahlgren et Clifford in Dahlgren, Clifford et Yeo, Fam. Monocot.: 175. 1985

Genera/species 1/2

Distribution Coastal eastern Australia.

Fossils Unknown.

Habit Bisexual, perennial giant herbs. Bulb consisting of large leaf bases rich in starch. Secondary bulbs developing from lower axillary shoot primordia following anthesis.

Vegetative anatomy Phellogen absent. Secondary lateral growth absent. Vascular bundles enclosed by fibres. Vessel elements? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Idioblasts with calciumoxalate as single, double or triple crystals. Styloids present. Calciumoxalate raphides absent from vegetative organs (present in tepals).

Trichomes Hairs?

Leaves Alternate (spiral), simple, entire, long, often linear, with ? ptyxis. Stipules absent; leaf sheath well developed, enclosing stem. Venation parallelodromous. Stomata paracytic; subsidiary cells with oblique divisions. Cuticular wax crystalloids as parallel platelets (Convallaria type). Mesophyll with lignified parenchyma between vascular bundles, with scattered tanniniferous cells and with calciumoxalate as single prismatic crystals. Leaf margin entire. Epidermis at apex of older leaves caducous, leaving a tuft of dry threads (sometimes dissolved into fibres).

Inflorescence Terminal, panicle or capitate thyrse. Peduncle (scape) very long. Flowers sometimes replaced by bulbils.

Flowers Actinomorphic to somewhat zygomorphic, large. Epigyny. Tepals 3+3, petaloid, partially caducous, connate at base. Septal nectaries well developed, opening around stylar base. Disc absent.

Androecium Stamens 3+3. Filaments linear or widened at base, free from each other, often adnate in lower part to tepals (epitepalous). Anthers peltate, pseudo-basifixed (centrifixed), non-versatile, tetrasporangiate, latrorse (to introrse?), longicidal (dehiscing by longitudinal slits); filament apex enclosed by tube formed by massive connective; anthers sometimes adnate to tepal bases. Tapetum secretory, outer layer with uninucleate cells, inner layer with multinucleate cells. Staminodia absent.

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

Gynoecium Pistil composed of three connate carpels. Ovary inferior, trilocular. Style single, simple, narrow, with stylar canal. Stigma triangular to punctate, papillate?, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules five to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument five to seven cell layers thick. Inner integument two cell layers thick. Parietal cell formed from archesporial cell. Parietal tissue approx. five cell layers thick. Hypostase present. Nucellar cap approx. two cell layers thick, formed by periclinal cell divisions in megasporangial epidermis. Megagametophyte monosporous, Polygonum type. Antipodal cells persistent, proliferating (up to five cells). Endosperm development ab initio helobial. Endosperm haustoria? Embryogenesis?

Fruit A lignified loculicidal capsule.

Seeds Aril absent. Seeds often winged. Seed coat testal-tegmic. Testa multilayered, multiplicative, with phlobaphene and degradation products of this substance. Phytomelan absent. Tegmen thick, with distinct cuticle. Perisperm not developed. Endosperm copious, with thin-walled cells rich in oils and aleurone (starch absent). Embryo straight, flattened, chlorophyll? Cotyledon one, wide, obtriangular. Cotyledon hyperphyll? Hypocotyl internode? Coleoptile present. Germination? Seedling with laterally compressed haustorium.

Cytology n = 17, 18, 22, 24. – Karyotype bimodal.

DNA

Phytochemistry Cyanidin and steroidal saponins present. Flavonols, ellagic acid, and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Doryanthes (2; coastal areas in southeastern Queensland and eastern New South Wales). Doryanthaceae are sometimes recovered as sister-group to Iridaceae (e.g. Rudall 2003).

HEMEROCALLIDACEAE R. Br.

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Brown, Prodr. Fl. Nov.-Holl.: 295. 27 Mar 1810 [’Hemerocallidae’]

Phormiaceae J. Agardh, Theoria Syst. Plant.: 7. Apr-Sep 1858; Dianellaceae Salisb., Gen. Plant.: 66. 15-30 Mai 1866 [’Dianelleae’]; Johnsoniaceae J. P. Lotsy, Vortr. Bot. Stammesgesch. 3: 731. 3-20 Sep 1911; Geitonoplesiaceae R. Dahlgren ex Conran in Telopea 6: 39. Nov 1994; Eccremidaceae Doweld, New Syllabus Plant Fam.: 942. Apr 2007

Genera/species 19–20/85–105

Distribution Europe, tropical East Africa, southern and southeastern Africa, Madagascar, the Mascarene Islands, the Seychelles, West and Central Asia eastwards to China and Japan, Southwest, South and Southeast Asia, Malesia, New Guinea, Australia, New Caledonia, New Zealand (including the Chatham Islands), Norfolk Island, Fiji, Polynesia including the Hawaiian Islands, the Andes.

Fossils Pollen of Phormium is described from the Late Miocene of New Zealand.

Habit Dioecious, perennial herbs or shrubs (in Corynotheca deciduous; in Geitonoplesium climbing). Phyllocladia present in Hensmania, Hodgsoniola, Johnsonia, and Stawellia. Roots often swollen, contractile, sometimes tuberous (sometimes fibrous; rarely stilt roots).

Vegetative anatomy Root endodermis with stout U-formed cell wall thickenings (not in Arnocrinum). Phellogen? Secondary lateral growth absent. Vessels present in roots (in Caesia, Geitonoplesium, Johnsonia, and Tricoryne 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 P2c type, with cuneate protein crystals. Nodes? Mucilage cells and mucilage canals absent. Tanniniferous cells present in at least some species. Calciumoxalate as cubes or crystal sand; raphides absent.

Trichomes Hairs absent.

Leaves Alternate (usually distichous, sometimes spiral), simple, entire, usually linear (sometimes semi-equitant, unifacial), with conduplicate to conduplicate-flat ptyxis, often unifacially keeled. Stipules absent; leaf sheath closed. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids as parallel platelets (Convallaria type) or non-orientated platelets or rodlets. Mesophyll often with calciumoxalate as druses or single prismatic crystals. Leaf margin serrate or entire.

Inflorescence Terminal, usually panicle, sometimes raceme- or spike-like, possibly consisting of one or two double helicoid cymes (in Tricoryne with umbel-like partial inflorescences; in Hensmania, Hodgsoniola, Johnsonia, and Stawellia simple raceme or spike; flowers in Herpolirion solitary). Floral prophylls (bracteoles) sometimes (in, e.g., Dianella and Hemerocallis) lateral.

Flowers Usually actinomorphic (in Hemerocallis and Phormium zygomorphic), sometimes large. Pedicel usually articulated. Usually (secondary) hypogyny (in Pasithea half epigyny). Tepals 3+3 (median outer tepal in Hemerocallis adaxial), petaloid, usually marcescent (often twisted), free or connate in lower part into a tube (in Arnocrinum and Hemerocallis connate in lower half). Septal nectaries infralocular (in Johnsonia and Tricoryne external at ovary base; absent in Hemerocallis and Phormium). Androecial nectaries present in Dianella. Disc absent.

Androecium Stamens usually 3+3 (in Hodgsonia three). Filaments often ornamented, usually free from each other (in Eccremis slightly connate at base), often adnate to tepal bases (in Eccremis three stamens hypogynous and three stamens slightly adnate to tepals [epitepalous]); if stamens three then filaments adnate at base to inner tepals, somewhat upcurved (in Simethis and Stypandra more or less hairy; filaments in, e.g., Dianella, Rhuacophila and Stypandra with swelling, struma). Anthers usually basifixed (sometimes dorsifixed to centrifixed or peltate), sometimes versatile, tetrasporangiate, latrorse or introrse (in Corynotheca also extrorse), usually longicidal (dehiscing by longitudinal slits; in Dianella etc. pseudoporicidal, dehiscing by apical pores and subsequently longicidal; anthers in Geitonoplesium and Stypandra coiled after anthesis; anthers in Arnocrinum connate and forming a tube around style), often twisted. Tapetum secretory, with binucleate cells (remains of degenerating protoplasts sometimes invading anther locule in Dianella). Staminodia usually absent (in Hodgsonia three).

Pollen grains Microsporogenesis usually simultaneous (in Hemerocallis successive). Pollen grains usually trichotomosulcate (in Hemerocallis monosulcate), shed as monads, bicellular at dispersal. Exine tectate (sometimes semitectate) or intectate, with columellate infratectum, usually perforate or microreticulate (in Hemerocallis and Phormium reticulate), tuberculate.

Gynoecium Pistil composed of three connate carpels (in Tricoryne almost free). Ovary usually (secondarily) superior (rarely semi-inferior), trilocular or unilocular. Style single, simple, filiform, usually terminal (in Tricoryne gynobasic). Stigma small, usually capitate or punctate (in Hemerocallis as a short hair tuft; in Pasithea trilobate), papillate?, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile (ovary trilocular; sometimes basal-axile or apical-axile) or parietal (ovary unilocular). Ovules one to c. 30 (to more than 50) per carpel, anatropous or campylotropous, ascending to pendulous, bitegmic, tenuinucellar (to weakly crassinucellar). Micropyle endostomal. Outer integument six or seven cell layers thick. Inner integument two to four cell layers thick. Parietal cell not formed (parietal tissue absent). Nucellar cap approx. two cell layers thick, sometimes formed by periclinal cell divisions starting in epidermal layer of megasporangium (absent in Hemerocallis). Hypostase absent. Megagametophyte monosporous, Polygonum type. Chalazal nucleus well developed. Synergids with a filiform apparatus (at least in some genera). Antipodal cells large, persistent. Endosperm development usually helobial (in Hemerocallis nuclear). Endosperm haustoria? Embryogenesis?

Fruit Usually a dry loculicidal capsule (in Eccremis fleshy, with loculicidal exocarp and septicidal endocarp; sometimes a berry; in Corynotheca a nut; in Tricoryne a schizocarp).

Seeds Seed often with strophiole or aril (from hilum; in Phormium winged). Elaiosome absent. Seed coat exotestal. Exotesta thick, with phytomelan layer on epidermal cell walls. Endotesta compressed. Tegmen collapsed. Perisperm not developed. Endosperm copious, with oil and aleurone (with usually little or no hemicellulose or starch). Embryo long or short, usually straight (in Arnocrinum curved), well differentiated, without chlorophyll. Cotyledon one, unifacial, with a prolonged closed and tubular sheath (in Caesia, Dianella and Tricoryne epigeal hyperphyll but not cotyledon photosynthesizing). Cotyledon hyperphyll elongate, assimilating, or compact, not assimilating. Epicotyl often longer. Hypocotyl internode short (sometimes absent). Coleoptile absent. Collar sometimes present. Radicula well developed, often branched, persistent. Germination?

Cytology n = 4, 8, 9, 11, 12, 14, 16, 22, 24, 38, 40, 42 – Chromosomes 0,8–17 µm long.

DNA The mitochondrial gene rpl2 is absent (lost). Loss of the 3’-rps12 intron is a synapomorphy of the clade [Hemerocallidoideae+Johnsonioideae].

Phytochemistry Flavonols (kaempferol, quercetin, in Hemerocallis), phenolic glycosides, steroidal saponins (at least in Pasithea), naphthoquinones (e.g. imbricatonol), chelidonic acid, and polyacetate derived arthroquinones present. Eicosanyl arachidate, chrysophanol, and nepodin present in roots. Dianellidin, dianellidone, imbricatonol, stypandrone, and stypandrol (hemerocallin, a toxic binaphthalene tetrol) present in, e.g., Dianella, Hemerocallis and Stypandra. Cyanidin present in Dianella. Ellagic acid not found.

Use Ornamental plants, medicinal plants, textile plants (Phormium tenax).

Systematics Hemerocallidaceae are probably sister to Xanthorrhoeaceae. However, Asphodelaceae have sometimes been recovered as sister to Hemerocallidaceae (e.g. McPherson & al. 2004; Chase & al. 2006). The loss of the 3’-rps12 intron is a character present in both Hemerocallidoideae and Johnsonioideae.

Phormioideae (J. Agardh) Thorne et Reveal in Bot. Rev. (Lancaster) 73: 80. 29 Jun 2007

10/30–50. Pasithea (1; P. caerulea; Chile); Phormium (2; New Zealand, Stewart Island, Norfolk Island, Chatham Islands), Agrostocrinum (1; A. scabrum; southwestern Western Australia), Geitonoplesium (1; G. cymosum; the Philippines, Central and East Malesia eastwards to New Guinea, eastern Queensland, eastern New South Wales, eastern Victoria, Norfolk Island, Lord Howe, New Caledonia, Fiji), Thelionema (3; southeasternmost Queensland, eastern New South Wales, Victoria, Tasmania), Rhuacophila (1; R. javanica; Malesia, New Caledonia, Fiji), Herpolirion (1; H. novae-zelandiae; eastern Victoria, Tasmania, New Zealand), Stypandra (1; S. glauca; southwestern Western Australia, southern South Australia, southeastern Queensland to Victoria, New Caledonia), Dianella (20–40; East Africa, Madagascar, East Asia to Japan, tropical Asia from India to Malesia, Australia, New Zealand, southwestern Pacific islands, the Hawaiian Islands, with their highest diversity in Australia), Eccremis (1; E. coarctata; the Andes). – East Africa, Madagascar, East and tropical Asia, Australia, Melanesia, New Zealand, Polynesia, the Andes, with their largest diversity in southwestern and eastern Australia. Leaves often bifacial basally and distally, being semi-ensiform and unifacial in middle just after diverging from sheath (Wurdack & Dorr 2009, etc.). – Pasithea, with entirely bifacial leaves, is sister to the remaining Phormioideae. The Andean Eccremis coarctata (sister to Dianella) has articulated pedicels, filaments connate at base, three filaments adnate to tepals, and a septicidal endocarp. Aloin cells (see Asphodelaceae) have been reported from Dianella.

[Hemerocallidoideae+Johnsonioideae]

Hemerocallidoideae (Bartl.) Lindl., Veg. Kingd.: 201, 205. Jan-Mai 1846 [‘Hemerocallideae’]

2/c 16. Hemerocallis (c 15; Central Europe, temperate Asia eastwards to China and Japan), Simethis (1; S. planifolia; southwestern Europe to southwestern Ireland and Italy, western Mediterranean). – Eurasia. – Hemerocallidoideae are sister-group to Johnsonioideae, according to the analyses by Wurdack & Dorr (2009).

Johnsonioideae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 233. 20 Jul 1943

7/38. Tricoryne (7; New Guinea, Australia), Corynotheca (6; Australia), Caesia (12; Northern, Western and Eastern Cape, Madagascar, New Guinea, Australia), Arnocrinum (3; southwestern Western Australia), Stawellia (2; southwestern Western Australia), Hensmania (3; southwestern Western Australia), Johnsonia (5; southwestern Western Australia). – South Africa, Madagascar, New Guinea, Australia, with their highest diversity in southwestern Western Australia. Aril present. – Tricoryne is sister to the remaining Johnsonioideae. Hodgsoniola (1; H. junciformis; southwestern Western Australia), sometimes included in Anthericaceae and possessing phylloclades, is possibly closely allied to the clade [Stawellia+[Johnsonia+Hensmania]].

Maximum likelihood bootstrap consensus tree of Hemerocallidaceae based on DNA sequence data (Wurdack & Dorr 2009).

HERRERIACEAE Kunth

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Kunth, Enum. Plant. 5: 290. 10-11 Jun 1850 [’Herrerieae’]

Genera/species 2/9

Distribution Madagascar, South America.

Fossils Unknown.

Habit Bisexual, perennial herbs or suffrutices, usually climbing or twining. Rhizome often tuberous. Climbing roots absent. Aerial stem spiny or absent.

Vegetative anatomy Roots fibrous; root exodermis uniseriate. Phellogen absent? Stem cortex with a cylinder of fibre bundles and small collateral vascular bundles; centre of stem with scattered bundles. Rhizome in Herreria with lignified secondary xylem and parenchymatous secondary phloem. Aerial stem at least in Herreria montevidensis with secondary lateral growth. Vessels present in roots and in Herreria also in stem. Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Mucilage cells and laticifers absent. Tanniniferous cells present. Calciumoxalate raphides frequent.

Trichomes Hairs absent?

Leaves Alternate (spiral), simple, entire, often linear, cladode-like, with supervolute (Herreria) ptyxis. Stipules and leaf sheath absent. Venation parallelodromous. Stomata anomocytic, in parallel rows on adaxial side (Herreria). Cuticular wax crystalloids as parallel platelets (Convallaria type). Mesophyll with cells containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal or axillary, panicle or raceme.

Flowers Actinomorphic. Pedicel not articulated. Hypogyny. Tepals 3+3, petaloid, persistent or caducous, free. Septal nectaries present in Herreria. Outer tepals in Herreriopsis with saccate tepal nectaries at base. Disc absent.

Androecium Stamens 3+3. Filaments free from each other and from tepals. Anthers basifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory (tapetal material slightly invading anther locule at dyad stage). Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, perforate, microreticulate or reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple, narrow. Stigma capitate or trilobate, type? Pistillodium absent.

Ovules Placentation intrusively axile. Ovules one to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument four cell layers thick. Inner integument two cell layers thick. Hypostase present. Parietal tissue? Nucellar cap present? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio helobial. Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule with partially persistent tepals.

Seeds Aril absent. Seeds flattened, spirally winged. Exotesta with phytomelan layer on outer epidermal cell walls, collapsed. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, with oils and aleurone (starch absent); endosperm cell walls without pits. Embryo small, straight, chlorophyll? Cotyledon one, without sheath. Cotyledon hyperphyll long, photosynthesizing. Hypocotyl internode? Coleoptile? Germination cryptocotylar?

Cytology n = 27 (Herreria) – Chromosomes 0.7–3.7 µm long, dimorphic (bimodal, with one large chromosome).

DNA The mitochondrial gene rpl2 absent.

Phytochemistry Flavonols, tannins, phenols, steroidal saponins (e.g. gitogenin in roots), and water-soluble steroids present. Chelidonic acid? Ellagic acid not found.

Use Unknown.

Systematics Herreria (8; Brazil and southwards in South America), Herreriopsis (1; H. elegans; Madagascar).

Herreriaceae is sister to Behnia (Behniaceae).

HYACINTHACEAE Batsch ex Borkh.

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Borkhausen, Bot. Wörterb. 1: 315. 1797 [’Hyacinthinae’]

Scillaceae Vest, Anleit. Stud. Bot.: 267, 284. 1818 [’Scilloideae’]; Hyacinthineae Link, Handbuch 1: 160. 4-11 Jul 1829 [’Hyacinthinae’]; Eucomidaceae Salisb., Gen. Plant.: 16. 15-31 Mai 1866 [’Eucomeae’]; Lachenaliaceae Salisb., Gen. Plant.: 20. 15-31 Mai 1866 [’Lachenaleae’]; Ornithogalaceae Salisb., Gen. Plant.: 33. 15-31 Mai 1866 [’Ornithogaleae’]

Genera/species 38–43/820–855

Distribution Europe, Africa, southwestern Asia and eastwards to Burma and Sri Lanka, southwestern South America (Chile and the Andes), with their largest diversity in South Africa, the Mediterranean and southwestern Asia.

Fossils Unknown.

Habit Usually bisexual (rarely polygamomonoecious), perennial herbs. Usually bulbous.

Vegetative anatomy Bulb usually with a membranous tunica; bulb scales free or more or less coalescent. Roots usually contractile; carbohydrate storing roots present in Ledebouria. Phellogen absent. Stem and leaves with large rhexigenetic lacunae. Secondary lateral growth absent. Vessels present in roots. Vessel elements with scalariform and/or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Mucilage cells abundant, often with calciumoxalate raphides.

Trichomes Hairs absent.

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, often linear or filiform, with ? ptyxis. Stipules absent; leaf sheath well developed, bulb scale with closed or open sheath. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids usually as non-orientated platelets (sometimes parallel platelets, Convallaria type). Mesophyll with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin entire.

Inflorescence Terminal, simple, raceme (rarely spike, head or branched raceme; flowers rarely solitary) on elongated peduncle (scape). Floral prophylls (bracteoles) usually absent.

Flowers Usually actinomorphic (in Lachenalia and Daubneya aurea zygomorphic with median outer tepal adaxial; inner tepals rarely strongly reduced). Pedicel usually not articulated (in Schizobasis articulated). Usually hypogyny (in Bowiea half epigyny). Tepals 3+3, petaloid, free or connate into a tubular, campanulate or urceolate perianth. Corona sometimes present. Septal nectaries usually present, of simple shape (rarely basally connate; absent in Autonoe; in some species of Ornithogalum also a stylar nectary). Disc absent.

Androecium Stamens usually 3+3 (in, e.g., some species of Ornithogalum three fertile stamens and three staminodia). Filaments filiform or broad and flat (sometimes with apical or basal appendages), usually free (sometimes connate at base; in Puschkinia forming a paracorolla), often adnate at base to tepals. Anthers usually dorsifixed, versatile?, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum secretory, with binucleate cells. Staminodia usually absent (sometimes three, extrastaminal).

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

Gynoecium Pistil composed of three connate carpels (sometimes syncarpously connate in lower part and paracarpously connate in upper part). Ovary usually superior (rarely semi-inferior), trilocular. Style single, simple (sometimes very short), with a stylar canal. Stigma punctate to somewhat trilobate, papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules two to c. 50 (in Barnardia one) per carpel, anatropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument three or more cell layers thick. Inner integument two cell layers thick. Obturator often present. Parietal cell formed from archesporial cell. Parietal tissue usually two to four cell layers thick. Nucellar cap often present. Megasporangial epidermis sometimes radially elongate. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, or tetrasporous, Scilla type [Hyacinthoides type, Endymion type], Adoxa type, DrusaI type, or Fritillaria type). Synergids with a filiform apparatus. Endosperm development usually helobial (sometimes nuclear). Endosperm haustoria? Embryogenesis asterad, caryophyllad or chenopodiad.

Fruit A loculicidal capsule, dry or fleshy.

Seeds Aril absent. Caruncular elaiosome sometimes present. Seed coat exotestal. Exotesta usually with phytomelan layer on epidermal cell walls (absent in some species of Hyacintheae). Mesotesta, endotesta and tegmen usually collapsed. Perisperm not developed. Endosperm copious, with oils, aleurone and hemicellulose (starch usually absent); endosperm cells with pitted walls. Embryo large, usually straight (sometimes curved), without chlorophyll. Cotyledon one. Cotyledon hyperphyll elongate and assimilating, or compact and non-assimilating. Hypocotyl internode absent. Coleoptile absent. Collar rhizoids sometimes present. Germination cryptocotylar? or phanerocotylar? Seedling foliage distichous.

Cytology x = 2–12 – Polyploidy and aneuploidy frequently occurring (even deca- and dodecaploidy). Chromosomes 1.2–18 µm long; chromosomes sometimes bimodal or trimodal.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols (kaempferol, quercetin), flavones, flavone-C-glycosides, flavonoid sulphates, homoisoflavanones, polyhydroxyalkaloids, cholestane glycosides (cardiotoxic bufodienolides [e.g. scilliroside] and cardenolides), steroidal saponins, cyanogenic compounds, salicylic acid, and chelidonic acid present. Tricine rare. Carbohydrates in bulb mainly stored as fructans (sometimes also as starch). Ellagic acid, proanthocyanidins and alkaloids not found.

Use Ornamental plants, medicinal plants, vegetables (Muscari).

Systematics Hyacinthaceae are sister to Themidaceae.

A plausible topology is [Oziroeoideae+[Urgineoideae+[Ornithogaloideae+Hyacinthoideae]]].

Oziroeoideae Speta in Phyton (Horn) 38: 51. 14 Aug 1998

1/5. Oziroe (5; southwestern South America). – Filaments at base connate and adnate to inner tepals. Seeds rounded. Embryo as long as seed. n = 15, 17. –Oziroe is sister to the remaining Hyacinthaceae.

[Urgineoideae+[Ornithogaloideae+Hyacinthoideae]]

Rhexigenetic lacunae present. Styloids present.

Urgineoideae Speta in Phyton (Horn) 38: 51. 14 Aug 1998

3–8?/115–120. Bowiea (1; B. volubilis; Uganda, Kenya and southwards to South Africa); Drimia (c 160?; the Mediterranean, Africa, Madagascar, India), Litanthus (5–10?; southern Africa; in Drimia?)?, Rhadamanthus (c 20; Namibia, Northern, Western and Eastern Cape, Free State; in Drimia?)?, Sagittanthera (2; eastern Eastern Cape; in Drimia?), Schizobasis (9; Ethiopia and southwards to South Africa; in Drimia?)?, Tenicroa (5; Namibia, Northern, Western and Eastern Cape; in Drimia?)?, Thuranthos (c 10; southern Africa, Zimbabwe, Zambia, Mozambique; in Drimia?)? – The Mediterranean, Africa, Madagascar, the Arabian Peninsula to Sri Lanka and Burma. Bracts with spurs (in Bowiea as small leaves). Seeds flattened and/or winged. Testa fragile, not tightly adherent to endosperm. n = 6, 7, 10 or more. Bufadienolides present. – Urgineoideae are sister-group to [Ornithogalum+Hyacinthoideae]. Bowiea is sister to the remaining Urgineoideae.

[Ornithogaloideae+Hyacinthoideae]

Ornithogaloideae Speta in Phyton (Horn) 38: 51. 14 Aug 1998

1/270–300. Ornithogalum (270–300; Europe, the Mediterranean, subtropical and southern Africa, Madagascar, southern Arabian Peninsula, Socotra, southwestern Asia and eastwards to India and Sri Lanka, with their largest diversity in South Africa). – Stamens sometimes three. Filaments sometimes flat, provided with appendages. Seeds flattened and/or short. Cell nuclei containing protein crystalloids. Cotyledon often photosynthesizing. n = 2–10 or more. Cardenolides present.

Hyacinthoideae Speta in Phyton (Horn) 38: 51. 14 Aug 1998

33/430–435. Leaves sometimes with pustulae or coloured spots. Megagametophyte varying. Seeds often rounded. Elaiosome sometimes present. Homoisoflavanones present.

Pseudoprospereae J. C. Manning et Goldblatt in Edinburgh J. Bot. 60: 557. 14 April 2004

1/1. Pseudoprospero (1; P. firmifolium; southern and eastern South Africa). – Prophylls present. Ovules two per carpel. One seed per locule. Cotyledon non-photosynthesizing. n = 9. – Pseudoprospero is probable sister to [Massonieae+Hyacintheae].

[Massonieae+Hyacintheae]

Massonieae Baker in Bot. J. Linn. Soc. London 11: 355. 17 Sep 1870

12/190–195. Merwilla (5; southern Africa to Zimbabwe); Eucomis (c 10; South Africa and northwards to Zimbabwe and Malawi); Spetaea (1; S. lachenaliiflora; Western Cape), Daubenya (8; Northern and Western Cape); Veltheimia (2; Northern, Western and Eastern Cape), Massonia (c 10; Namibia, South Africa, Lesotho), Lachenalia (c 110; Namibia, Northern, Western and Eastern Cape, Free State), Namophila (1; N. urotepala; southern Namibia); Schizocarphus (1; S. nervosus; tropical East to southern Africa), Ledebouria (15–20; tropical and southern Africa, Madagascar, southern India), Drimiopsis (c 20; tropical and southern Africa), Resnova (c 10; southern Africa). – Tropical and southern Africa, Madagascar, southern India. Floral prophylls (bracteoles) sometimes present. Flowers sometimes zygomorphic. Filament tube present in some species of Daubenya. Ovary and style usually sulcate. Style with three canals. Ovules two to numerous per carpel. Elaiosome sometimes present. Cotyledon usually non-photosynthesizing. n = 5–10 or more. – Merwilla seems to be sister to the remaining Massonieae.

Hyacintheae Dumort., Fl. Belg.: 141. 1827

21/c 240. Barnardia (1; B. japonica: eastern China, the Korean Peninsula, Japan, Russian Far East); Tractema (6; western Europe, western Mediterranean, northwesternmost Africa), Oncostema (c 10; Iberian Peninsula, western Mediterranean, northwesternmost Africa), Autonoe (6; Macaronesia, the Mediterranean), Hyacinthoides (3–4; western Europe to North Africa); Brimeura (3; western Mediterranean), Hyacinthella (18; southeastern Europe, southwestern Asia), Alrawia (2; Iran, Iraq), Prospero (c 25; western, southern and southeastern Europe, theMediterranean, northernmost Africa, Turkey, Caucasus), Puschkinia (3–4; Turkey, Lebanon, Syria, Caucasus), Othocallis (c 20; Europe, the Mediterranean, western Asia, the Middle East to Iran), Fessia (10; Iran, Pakistan, Central Asia), Pfosseria (1; P. bithynica; eastern Balkan Peninsula, western Turkey), Zagrosia (1; Z. persica; southestern Turkey, Iraq, western Iran), Hyacinthus (3; western and Central Asia); Chouardia (2; the Balkan Peninsula), Nectaroscilla (1–2; eastern Mediterranean), Schnarfia (2; southern Balkan Peninsula), Muscari (c 50; Europe, the Mediterranean, western and southwestern Asia), Bellevalia (45–50; the Mediterranean and eastwards to Iran and northern Afghanistan), Scilla (c 30; Europe, Macaronesia, the Mediterranean, North Africa, West Asia). – Europe to southwestern Asia, North Africa, East Asia. Bract sometimes absent. Prophylls fairly frequently present. Style with a single papillate canal. Ovules two to eight (to numerous) per carpel. Outer integument four or five cell layers thick. Parietal tissue two or three cell layers thick. Antipodal cells large. Elaiosome sometimes present. Phytomelan sometimes absent. Cotyledon often photosynthesizing. n = 4–8 or more. – Barnardia japonica seems to be sister to all other Hyacintheae. ‘Barnardianumidica (the Balearic Islands, Algeria, Tunisia, Libya) is obviously not closely related to Barnardia japonica (Ali & al. 2012).

Cladogram (simplified) of Hyacinthaceae based on DNA sequence data (Pfosser & Speta 1999; with the addition of Pseudoprospereae).

HYPOXIDACEAE R. Br.

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Brown in M. Flinders, Voy. Terra Austral. 2: 576. 19 Jul 1814 [’Hypoxideae’], nom. cons.

Hypoxidales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 9. Sep-Oct 1835 [‘Hypoxideae’]

Genera/species c 6/100–220?

Distribution Africa, the Mascarene Islands, the Seychelles, tropical and subtropical regions in Asia northwards to eastern Himalaya, southwestern China and Japan, New Guinea, Australia, Tasmania, New Zealand, eastern USA to the West Indies and South America southwards to Uruguay, with the largest diversity in South Africa.

Fossils Unknown.

Habit Usually bisexual (in Curculigo rarely unisexual), perennial herbs. Tuberous rhizome or corm.

Vegetative anatomy Velamen present or absent. Roots often contractile. Dimorphic root hypodermis absent. Phellogen absent. Rhizome and stem nodules with inner parenchymatous zone containing irregularly running vascular bundles and fibrous tissue, surrounded by starchy parenchyma usually with secretory cavities (mucilage canals). 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 P2c type, with cuneate protein crystals. Silica bodies absent. Calciumoxalate raphides usually present.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, often dichotomously branched and with multicellular base, sometimes stellate or lepidote.

Leaves Alternate (usually tristichous), simple, entire, often linear, sometimes plicate or unifacial and subulate in transverse section (in Curculigo, Hypoxidia and Molineria differentiated into pseudopetiole and pseudolamina), with plicate to conduplicate-plicate (sometimes conduplicate-flat) ptyxis. Stipules absent; leaf sheath open or closed, persistent. Venation parallelodromous (sometimes pinnate-parallel), with distinct main veins. Stomata usually paracytic (rarely tetracytic), with oblique or parallel cell divisions. Cuticular waxes absent. Secretory cavities (mucilage canals) present above vascular bundles in some genera (in Empodium in mesophyll). Mesophyll with calciumoxalate raphides. Leaf margin entire.

Inflorescence Axillary, often on long peduncle (scape), corymbose, spike- or umbel-like cymose (flowers sometimes solitary).

Flowers Actinomorphic. Epigyny. Tepals usually 3+3 (in Hypoxis and Spiloxene rarely 2+2), petaloid, often with valvate aestivation, persistent, usually free (in Hypoxidia, Pauridia and Saniellia connate into a tube). Septal nectaries absent. Disc absent.

Androecium Stamens usually 3+3 (rarely 2+2; in Pauridia three antepetalous stamens and three staminodia adnate to style into a gynostemium). Filaments short, filiform or subulate, free, often adnate to lower part of tepals (epitepalous). Anthers basifixed, centrifixed or slightly dorsifixed, occasionally somewhat versatile, tetrasporangiate, usually introrse or latrorse (sometimes extrorse), longicidal (dehiscing by longitudinal slits); connective rarely somewhat prolonged at apex. Tapetum secretory or amoeboid-periplasmodial, with uni- to multinucleate cells. Staminodia usually absent (in Pauridia three intrastaminal staminodia).

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (sometimes trisulcate or inaperturate, in Pauridia disulcate), shed as monads, bicellular at disperal. Exine tectate or semitectate, with columellate infratectum, perforate or finely reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary inferior, usually trilocular (in Empodium unilocular; in Hypoxidia and some species of Spiloxene trilocular in lower part and unilocular in upper part); ovary often with rostrum (small apical elongation). Style single, simple, stout, or stylodia three, more or less connate; with a stylar canal. Stigma trilobate or consisting of three vertical furrows along upper part of style, papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation usually axile (in Empodium parietal). Ovules few to numerous per carpel, usually anatropous (in Empodium and Pauridia possibly hemianatropous to campylotropous), apotropous, bitegmic, pseudocrassinucellar. Micropyle bistomal (sometimes Z-shaped, zig-zag), often prolonged. Outer integument two cell layers thick. Inner integument two cell layers thick. Parietal cell sometimes formed (from archesporial cell), degenerating (parietal tissue absent). Hypostase present. Nucellar cap formed by periclinal cell divisions in megasporangial epidermis, two or three cell layers thick, or absent. Megagametophyte usually monosporous, Polygonum type (in Empodium sometimes disporous, Allium type), with chalazal constriction. Synergids sometimes with a filiform apparatus. Antipodal cells ephemeral or persistent. Endosperm development usually helobial (in Pauridia and sometimes in Spiloxene nuclear). Endosperm haustorium chalazal or absent. Embryogenesis onagrad, solanad or asterad.

Fruit Usually a loculicidal capsule or a pyxidium (capsule sometimes denticidal, poricidal or irregularly dehiscing – with ovary beak – or baccate), usually with persistent parts of tepals.

Seeds Raphe prominent. Strophiole present in Curculigo and Empodium in association with hilum. Seed coat exotestal. Exotesta with phytomelan layer on epidermal cell walls, often palisade. Endotesta? Exotegmen? Endotegmen sometimes persistent. Perisperm not developed. Endosperm copious, with oils and aleurone (starch absent). Embryo small, straight, often little differentiated, chlorophyll? Cotyledon one, non-photosynthesizing, with reduced open leaf sheath and with apex transformed into haustorium. Cotyledon hyperphyll compact, non-assimilating. Hypocotyl internode and mesocotyl absent. Coleoptile present. Germination?

Cytology n = 6–9, 11, 16, 18, 19, 27, 36, 48 or more – Polyploidy occurring at least in Hypoxis. Chromosomes 2–5 µm long. Agamospermy present inHypoxis.

DNA

Phytochemistry Flavonols (quercetin) and chelidonic acid present. Ellagic acid, proanthocyanidins, alkaloids, steroidal saponins, and cyanogenic compounds not found. Ferulic acid absent from cell walls.

Use Ornamental plants.

Systematics Hypoxis (agamospermy present, with perhaps 50–100 microspecies; tropical and southern Africa, East and Southeast Asia, tropical to temperate America, with their highest diversity in South Africa), Empodium (9–10; South Africa, Swaziland, Lesotho), Pauridia (c 55; Namibia, Northern, Western and Eastern Cape, KwaZulu-Natal, Lesotho, Australia, Tasmania, New Zealand), ‘Curculigo’ (c 20; tropical regions on both hemispheres; polyphyletic), ‘Molineria’ (5–7; tropical Asia; paraphyletic; incl. Curculigo pro parte), Hypoxidia (2; the Seychelles).

Hypoxidaceae are probably sister-group to Lanaria (Lanariaceae) (or, possibly, to Asteliaceae).

Some analyses based on morphology indicate a close relationship between Hypoxidaceae and Orchidaceae.

Cladogram of Hypoxidaceae based on DNA sequence data (Kocyan & al. 2011).

IRIDACEAE Juss.

( Back to Iridales )

de Jussieu, Gen. Plant.: 57. 4 Aug 1789 [’Irides’], nom. cons.

Crocaceae Vest, Anleit. Stud. Bot.: 266, 283. 1818 [’Crocoideae’]; Ixiaceae Horan., Prim. Lin. Syst. Nat.: 51. 2 Nov 1834 [’Jxiaceae. (Jrideae)’]; Ixiales Lindl., Key Bot.: 70. 15-30 Sep 1835; Galaxiaceae Raf., New Fl. N. Amer. 1: 72. Dec 1836 [’Galaxidia’]; Gladiolaceae Raf., Fl. Tellur. 4: 34. med 1838 [’Gladiolina’]; Geosiridaceae Jonker in Receuil Trav. Bot. Néerl. 36 [Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 60]: 477. seors. impr. 18 Mai 1939, nom. cons.; Hewardiaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 234. 20 Jul 1943, nom. illeg.; Iridineae Engl., Syllabus, ed. 2: 94. Mai 1898; Isophysidaceae (Hutch.) F. A. Barkley in Revista Fac. Nac. Agron. Medellin Univ. Antioquia 8: 152. Sep 1948

Genera/species c 65/1.920–1.925

Distribution Nearly cosmopolitan, with their largest diversity in southern Africa, eastern Mediterranean, southwestern Asia and western South America.

Fossils Uncertain. Fossil pollen grains, Liliacidites goldblattii, resembling those in Isophysis (and Doryanthes), were described from the latest Campanian to earliest Maastrichtian Vilui Basin sediments in Siberia.

Habit Bisexual, usually perennial (some species of Sisyrinchium annual) herbs (Klattia, Nivenia and Witsenia are suffruticose with a more or less lignified stem base). Usually with a rhizome (often tuberous) or a tunicated corm (with basal innovation; rarely bulb). Geosiris aphylla in Madagascar is an achlorophyllous mycotrophic holoparasite with scale-like membranous leaves.

Vegetative anatomy Roots with mycorrhiza; often contractile; root stele central, surrounded by endodermis often with lignified cell walls; root hairs absent; root hypodermis dimorphic; root nodules or fleshy roots present in some genera. Rhizome sometimes with lignified endodermis-like layer. Phellogen? Stem parenchyma often starchy. Secondary lateral growth anomalous (Klattia, Nivenia, Witsenia) or absent. Vessels present in roots (in some genera also in stem and leaves). Vessel elements with scalariform or simple perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids usually P2c type, with cuneate protein crystals (rarely P2ccl type, with cuneate and several additional loosely packed protein crystals). Secretory cavities containing mucilage present. Tanniniferous cells present in many genera. Idioblasts with calciumoxalate as styloids or single prismatic crystals usually frequent (raphides absent).

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

Leaves Alternate (distichous), simple, entire, often linear or equitant, sometimes terete and filiform, unifacial, isobifacial or heterobifacial, with ad-abaxially plicate or conduplicate (sometimes conduplicate-plicate) ptyxis. Stipules absent; leaf sheath open or closed (ligule present in Geissorhiza). Venation parallelodromous; midvein often distinct (sometimes thickened). Stomata usually anomocytic (subsidiary cells in Diplarrhena and some species of Moraea formed by oblique cell divisions). Cuticular wax crystalloids as parallel platelets, Convallaria type. Secretory cavities with mucilage. Mesophyll with calciumoxalate as single prismatic crystals. Leaf margin entire.

Inflorescence Terminal, either thyrse, panicle or cyme composed of one or more umbel-like, corymbose or reduced rhipidia, or spike-like with often unilaterally rotated flowers (flowers often solitary). Bracts of various shape, often spathe-like. Extrafloral nectaries rarely present on bracts.

Flowers Actinomorphic or zygomorphic, often large. Some species with hypanthium. Usually epigyny (in Isophysis hypogyny). Tepals 3+3, petaloid, often aristate, free (Isophysis, Aristea, Geosiris, most Iridoideae), or connate into an infundibuliform or tubular perianth; perianth in zygomorphic flowers often bilabiate, with adaxial tepal usually largest (often dome-shaped) and three abaxial tepals smallest and patterned with nectar guides. Septal nectaries usually present, with antepetalous orifices at stylar base (in Crocoideae), or tepal nectaries inserted at tepal bases (in most Iridoideae and Aristea spiralis); tepal nectaries present as nectariferous or oil-secreting glandular hairs, elaiophores, usually unicellular and inserted at inner tepals; androecial (on filaments) or gynoecial nectaries often present in, e.g., Iris (nectaries absent in Isophysis, Geosiris and most species of Aristea). Disc absent.

Androecium Stamens usually three (in Diplarrhena two: outer abaxial stamen absent), antesepalous (in some genera asymmetrically placed), representing outer staminal whorl (inner whorl absent). Filaments coarse to filiform, usually free (sometimes more or less connate), often adnate to tepal bases or to perianth tube when present. Anthers usually basifixed or subbasifixed (sometimes centrifixed), sometimes versatile, tetrasporangiate, usually extrorse (rarely latrorse, with wide connective, or introrse), usually longicidal (dehiscing by longitudinal slits; in Cobana poricidal, dehiscing with apical pores). Tapetum secretory, with usually binucleate (sometimes multinucleate) cells. Staminodia absent.

Pollen grains Microsporogenesis usually simultaneous (in at least Geosiris successive). Pollen grains usually monosulcate (rarely disulcate, trisulcate, trichotomosulcate, zonosulculate, dizonosulculate, spiraperturate, or inaperturate), sometimes operculate, shed as monads, usually bicellular at dispersal. Exine usually tectate or semitectate (in Diplarrhena and Patersonia intectate), with columellate infratectum, usually reticulate or microreticulate (sometimes perforate to microscabrate; rarely areolate or rugulate).

Gynoecium Pistil composed of three connate carpels; median carpel abaxial. Ovary usually inferior (in Isophysis superior), usually trilocular (in Hermodactylus unilocular). Style single, filiform, usually trilobate, often further branched or petaloid (in Zygotritonia entire), with thin to wide (rarely hairy or fimbriate) branches, with stylar canal; stylar branches often widened at apex, in Iridoideae longitudinally folded, often flattened or thickened and with paired apical appendages, in Iris, Moraea and some other Irideae tangentially flattened, wide, petaloid, with two apical appendages and an apical stigma. Stigma papillate (with unicellular bifid papillae), Dry type; stigmatic branches adaxial on stylar lobes, inserted at apex, along margins or on surface of stylar branches. Pistillodium absent.

Ovules Placentation usually axile (in Hermodactylis parietal). Ovules one to c. 50 per carpel, anatropous or campylotropous, bitegmic, crassinucellar or tenuinucellar. Micropyle endostomal or exostomal. Outer integument usually four to six cell layers thick (in Aristea, Klattia, Nivenia and Witsenia two or three cell layers thick). Inner integument two cell layers thick. Parietal cell formed from archesporial cell. Parietal tissue one or two cell layers thick. Hypostase formed in Ixioideae from megasporangium at chalazal end of megagametophyte and surrounding antipodal cells. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells in some genera proliferating. Endosperm development usually nuclear (in Geosiris and Isophysis helobial). Endosperm haustoria? Embryogenesis asterad or caryophyllad.

Fruit Usually a loculicidal capsule (rarely indehiscent), hard (sometimes lignified), usually xerochastic (rarely hygrochastic).

Seeds Aril usually absent (present in some species of Iris). Elaiosome (arising from chalaza) present in Hermodactylus and Patersonia (in Patersonia also from raphe). Seed coat testal-tegmic (rarely winged). Exotesta usually with phlobaphene (absent in Klattia, Nivenia, and Witsenia), without phytomelan. Endotesta often with lipids. Tegmen often partially collapsed. Exotegmen and endotegmen usually pigmented. Perisperm not developed. Endosperm copious, hard, thick-walled, with oils, proteins and aleurone (rarely starch), with hemicellulose in pitted cell walls. Embryo small to fairly large, straight, usually without chlorophyll. Cotyledon one, terminal, coleoptile-like, usually not photosynthesizing (in, e.g., Sisyrinchium photosynthesizing; in, e.g., Tigridia with ligule or coleoptile). Cotyledon hyperphyll elongate or compact, often assimilating. Hypocotyl internode short or absent. Mesocotyl absent. Plumule lateral. Germination phanerocotylar.

Cytology n = (3–)5–17, 19, 20 – Polyploidy and aneuploidy occurring (in, e.g., Crocus, Iris, Moraea, and Sisyrinchium).

DNA

Phytochemistry Flavonols (kaempferol, quercetin etc.), flavone-C-glycosides, isoflavones (irigenin, tectorigenin, irisolon, and numerous others), amentoflavone (a biflavone present in Isophysis and Patersonia), cyanidin, tannins, alkaloids (e.g. homeridin), steroidal saponins, naphthoquinones (plumbagin), anthraquinones, polyacetate derived arthroquinones (in Gladiolus and Libertia), quinonoid pigments, mangiferin (a glycosylic xanthone), crocein (a yellow carotenoid in Crocus), hydroxycinnamic acid, chelidonic acid, and meta-carboxysubstituted aromatic amino- and γ-glutamylic peptides present. Tricine rare. Ellagic acid not found. Carbohydrates (e.g. saccharose, fructan and starch) stored in corms and rhizomes.

Use Ornamental plants, flavours (styles and stigmas from Crocus sativus, containing β- and γ-carotene, lycopin, zeaxanthin, crocin, safranal, etc.), perfumes (rhizome of Iris germanica), medicinal plants.

Systematics Iridaceae are sister-group to the remaining Iridales, comprising Xeronemataceae to Ruscaceae. Isophysis (Tasmania) is sister to all other Iridaceae.

Isophysidoideae (Hutch.) Takht. ex Thorne et Reveal in Bot. Rev. (Lancaster) 73: 79. 29 Jun 2007

1/1. Isophysis (1; I. tasmanica; Tasmania). – Rhizomatous herb. Root vessel elements with scalariform perforation plates. Flower solitary, spathaceous, with a pair of ’bracts’ subtending the single flower (perhaps corresponding to a reduced rhipidium). Hypogyny. Tepals free. Septal nectaries absent. Pistil composed of three connate carpels. Style shortly trifid, branches alternating with anthers. Stigmatic surfaces adaxial. Seedling unknown. n = ? Amentoflavone (a biflavonoid) present. – Isophysis tasmanica is sister to all other Iridaceae.

[Iridoideae+[Patersonioideae+[Geosiridoideae+[Aristeoideae+[Nivenioideae+Crocoideae]]]]]

Root vessel elements with usually simple perforation plates. Inflorescence usually rhipidium or spike, with cymose partial inflorescences with flowers successively arising from axils of prophylls. Epigyny. Septal nectaries, tepal nectaries or oil glands often present. Pollen grains sometimes operculate and sulcus often with two exine bands. Endosperm development usually nuclear. Xanthone (mangiferin) present.

Iridoideae Eaton, Bot. Dict., ed. 4: 28. Apr-Mai 1836 [‘Irideae’]

c 31/c 800. Diplarrhena (2; southeastern New South Wales, southern Victoria, Tasmania); Libertia (c 15; New Guinea, eastern New South Wales, Victoria, Tasmania, New Zealand, the Andes in Chile), Orthrosanthus (10; southwestern Western Australia, Central America, South America), Tapeinia (1; T. pumila; southern Chile, southern Argentina), Solenomelus (2; Chile, Argentina), Sisyrinchium (c 140; North America, Mexico, Central America, South America), Olsynium (14; western North America, southern Chile, southern Argentina); Neomarica (10–12; tropical America; in Trimezia?), Trimezia (c 40; tropical America; incl. Neomarica?), Pseudiris (1; P. speciosa; Chapada Diamantina in Brazil); Cypella (c 20; Mexico and southwards to Argentina), Hesperoxiphion (5; the Andes in Colombia, Peru and Bolivia), Cipura (9; tropical America), Calydorea (14; southern Chile, southern Argentina), Eleutherine (2; tropical America), Onira (1; O. unguiculata; Brazil), Gelasine (4; subtropical South America), Ennealophus (5; South America), Herbertia (5; southern Chile, southern Argentina, one species, H. lahue, to southern United States), Kelissa (1; K. brasiliensis; southern Brazil), Mastigostyla (16; Peru, Bolivia, Argentina), Cardenanthus (8; South America), Nemastylis (5; southern United States, Mexico, Central America), Ainea (1; A. conzattii; Mexico), Alophia (5; tropical and subtropical America), Tigridia (c 45; Central and South America), Cobana (1; C. guatemalensis; Guatemala, Honduras), Bobartia (15; Western and Eastern Cape), Iris (c 280; temperate regions on the Northern Hemisphere), Ferraria (13; southern tropical Africa to South Africa), Moraea (c 200; the Mediterranean, Africa, southwestern Asia, with their highest diversity in Western Cape). – Temperate regions on the Northern Hemisphere, Africa, Australia, Tasmania, New Zealand, North America to southern South America, with their largest diversity in South Africa and South America. Rhipidia simple. Habitus varying. Tepal nectaries usually present (in Diplarrhena septal nectaries). Oil glands or oil hairs often present. Anther endothecium with spiral thickenings. Stylar branches long and tubular. γ-glutamyl peptides and metacarboxy aminoacids present. – Five well supported main clades have been recognized in Iridoideae (Reeves & al. 2001): Diplarrhena (Australia), the sister-group to the remaining Iridoideae; Irideae; Sisyrinchieae; Tigridieae; and Mariceae.

[Patersonioideae+[Geosiridoideae+[Aristeoideae+[Nivenioideae+Crocoideae]]]]

Rhipidia two fused, each unit with two to numerous flowers; tepals connate; anther endothecium with base-plate or U-shaped thickenings; extra codon often present in gene rps4.

Patersonioideae Goldblatt in P. Goldblatt et J. C. Manning, Iris Fam.: 93. 12 Nov 2008

1/21. Patersonia (c 20; Sumatra, northern Borneo, New Guinea, Australia, Tasmania, New Caledonia). – Stem rhizomatous and often lignified. Secondary lateral growth present. Root vessel elements often with scalariform perforation plates. Inner tepals sometimes reduced to scales or absent. Septal nectaries absent. Filaments connate. Pollen grains spheroidal, inaperturate. Exine intectate. Embryo small. n = 11, 21. Two extra codons present in gene rps4. Amentoflavone (a biflavonoid) present.

[Geosiridoideae+[Aristeoideae+[Nivenioideae+Crocoideae]]]

Geosiridoideae Goldblatt et J. C. Manning, Iris Fam.: 95. 12 Nov 2008

1/2. Geosiris (1–2; G. aphylla, G. albiflora; Madagascar, Mayotte in the Comoro Islands). –Achlorophyllous root mycoparasitic herb. Leaves heterobifacial, reduced to scales. Flowers sessile. Tepals connate only at base. Septal nectaries absent. Microsporogenesis successive. Endosperm development helobial. Seeds extremely small. Endosperm with starch. n = ?

[Aristeoideae+[Nivenioideae+Crocoideae]]

Aristeoideae Vines, Stud. Text-book Bot. 2: 566. Mar 1895 [‘Aristinae’]

1/c 55. Aristea (c 55; South Africa to Senegal and Ethiopia, Madagascar, with their highest diversity in the Cape Provinces). – Root vessel elements with scalariform or simple perforation plates. Tepals connate only at base. Septal nectaries absent. Pollen morphology highly varying. Embryo small. n = 16. Plumbagin (a naphthoquinone) present.

[Nivenioideae+Crocoideae]

Flowers persistent, long-lived.

Nivenioideae Schulze ex Goldblatt in Ann. Missouri Bot. Gard. 77: 621. 16 Nov 1990

3/14. Nivenia (11; Western Cape), Klattia (3; Western Cape), Witsenia (1; W. maura; Western Cape). – Western Cape. Stem woody, with pronounced secondary lateral growth. Inflorescence biseriate, consisting of two terminal monochasia each with one or two flowers and a single bract. Tepals connate into a long tube. Pollen grains sometimes with encircling sulcus. Heterostyly present in Nivenia. Seeds peltate (tangentially flattened). Testa two or three cell layers thick. Exotesta transparent, not thickened. Exotegmen well developed. n = 16.

Crocoideae Burnett, Outlines Bot.: 451. Feb 1835 [’Crocidae’]

28/1.025–1.030. Watsonia (c 50; southern Africa, with their largest diversity in Western Cape), Thereianthus (8; Western Cape), Micranthus (3; Western Cape), Pillansia (1; P. templemannii; Caledon District in Western Cape), Lapeirousia (42; South Africa to Nigeria and Ethiopia, with their highest diversity in Northern and Western Cape), Savannosiphon (1; S. euryphyllus; Congo and Tanzania to Zambia, Malawi and northern Mozambique); Tritoniopsis (24; Western and Eastern Cape), Babiana (c 90; southern Africa, southern Zimbabwe, with their largest diversity in Northern and Western Cape), Cyanixia (1; C. socotrana; Socotra), Zygotritonia (4; western tropical Africa and eastwards to Ethiopia, Tanzania, Zambia and Socotra), Gladiolus (c 260; Europe, the Mediterranean, mountains in tropical Africa, Madagascar, southwestern Asia, with their highest diversity in Western Cape and the Drakensberg), Radinosiphon (1–3; R. leptostachya; northeastern South Africa to southern Tanzania), Xenoscapa (2; southern Namibia, Northern, Western and Eastern Cape), Geissorhiza (c 85; Northern, Western and Eastern Cape), Hesperantha (82; South Africa to Ethiopia, with their largest diversity in Western Cape and the Drakensberg), Melasphaerula (1; M. ramosa; southern Namibia, Northern and Western Cape), Romulea (c 90; the Canary Islands, the Mediterranean, Africa, Socotra, with their highest diversity in the Cape Provinces), Syringodea (8; southern Africa, especially Northern and Western Cape), Crocus (c 80; central and southern Europe, the Mediterranean, North Africa, southwestern Asia and eastwards to Central Asia and western China), Freesia (16; South Africa to Sudan, with their largest diversity in Western and Eastern Cape), Dierama (44; South Africa to Ethiopia), Ixia (c 65; Western and Eastern Cape), Sparaxis (15; Northern and Western Cape), Tritonia (28; South Africa to Tanzania), Duthiastrum (1; D. linifolium; northern Karroo in Northern Cape), Crocosmia (9; tropical and southern Africa, Madagascar), Devia (1; D. xeromorpha; Roggeveld Escarpment in Northern and Western Cape), Chasmanthe (3; Western and Eastern Cape). – Europe, the Mediterranean, Africa, Madagascar, Socotra, southwestern Asia to western China. Corm usually present. Root vessel elements with simple perforation plates. Leaves usually with pseudo-midvein (absent in Pillansia). Leaf sheath closed. Inflorescence spike-like or consisting of paired rhipidia usually with a single flower. Flowers actinomorphic or zygomorphic, often sessile. Tepals usually connate. Septal nectaries present or absent. Anther endothecium with spiral thickenings. Pollen grains with aperture having one or two longitudinal bands forming an operculum, sometimes zonaperturate. Exine tectate, perforate-scabrate. Ovules campylotropous. Chalazal hypostase prominent. Exotesta sometimes exfoliating. n = 9–17. – Two main clades are identified within Crocoideae, i.e. Ixieae and Watsonieae.

Cladogram (simplified) of Iridaceae based on DNA sequence data (Reeves & al. 2001).

IXIOLIRIACEAE (Pax) Nakai

( Back to Iridales )

Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 234. 20 Jul 1943

Genera/species 1/3

Distribution Egypt, Israel, eastern Turkey to Central Asia.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome a corm surrounded by a tunica.

Vegetative anatomy Root exodermis not dimorphic. 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 P2c type, with cuneate protein crystals. Nodes? Mucilage cells abundant. Calciumoxalate raphides frequent.

Trichomes Hairs bicellular to sexacellular, uniseriate.

Leaves Alternate (spiral), simple, entire, linear, with ? ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous, with unequally sized vascular bundles. Stomata anomocytic. Cuticular wax crystalloids? Mesophyll with calciumoxalate raphides. Leaf margin entire. Leaf apex short, subulate.

Inflorescence Terminal, leafy thyrse or almost umbel-like, sometimes botryoid or corymb with helicoid partial inflorescences (bostryces). Peduncle with sclerenchymatous cylinder; some vascular bundles of inflorescence axis arranged in a cylinder and enclosing remaining scattered bundles. Floral prophyll (bracteole) often lateral.

Flowers Actinomorphic. Epigyny. Tepals 3+3, petaloid, thin, connate at base; outer tepals with somewhat prolonged apex arising on abaxial side at a point immediately below apex. Septal nectaries present. Disc absent.

Androecium Stamens 3+3. Filaments free from each other, adnate in lower part (at base?) to tepals (epitepalous). Anthers basifixed to centrifixed, non-versatile, tetrasporangiate, introrse to latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate cells. Staminodia absent.

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

Gynoecium Pistil composed of three connate carpels. Ovary inferior, trilocular. Style single, simple, narrow. Stigma trilobate, papillate?, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules c. 15 to more than 50 per carpel, anatropous, ascending, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal cell formed from archesporial cell. Hypostase? Megagametophyte monosporous, Polygonum type. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule, apically dehiscing.

Seeds Aril absent. Seed coat testal. Exotesta with phytomelan layer on epidermal cell walls. Several testal layers with thin and reddish-brown cell walls. Mesotesta and endotesta with flat, empty cells. Tegmen crushed, membranous. Perisperm not developed. Endosperm copious, with thin and pitted cell walls, with oils and aleurone (starch almost absent, present in cells adjacent to embryo). Embryo large, straight, well differentiated, without chlorophyll. Cells surrounding embryo starchy. Cotyledon one. Cotyledon hyperphyll? Hypocotyl internode? Coleoptile? Germination?

Cytology n = 12

DNA

Phytochemistry Very insufficiently known. Chelidonic acid present. Steroidal saponins? Alkaloids not found.

Use Ornamental plants.

Systematics Ixiolirion (3; eastern Egypt, Israel, eastern Turkey and eastwards to Kashmir and Central Asia).

Ixiolirion is sister to Tecophilaeaceae.

LANARIACEAE H. Huber ex R. Dahlgren et A. E. van Wyk

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Dahlgren et A. E. van Wyk in Monogr. Syst. Bot. Missouri Bot. Gard. 25: 71. Jun 1988

Genera/species 1/1

Distribution The Cape Provinces in South Africa.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome vertical.

Vegetative anatomy Roots fibrous. 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 P2ccps type, with a single large loosely-packed polygonal protein crystal and many cuneate ones, and with starch grains. Nodes? Calciumoxalate raphides absent; cuboidal styloids sometimes present.

Trichomes Hairs unicellular, usually dendritic, long, especially on inflorescences and abaxial side of flowers.

Leaves Alternate (spiral or distichous), simple, entire, linear, with ? ptyxis. Stipules absent; leaf sheath closed? Venation parallelodromous. Stomata paracytic. Cuticular wax crystalloids? Epidermal cells associated with vascular strands and leaf margins often with lignified walls. Leaf margin entire.

Inflorescence Terminal, corymbose panicle with short branches. Inflorescence covered with dendritic hairs.

Flowers Actinomorphic. Epigyny to slightly half epigyny. Tepals 3+3, petaloid, connate in lower half into a tube. Abaxial side of tepals covered with dendritic hairs. Adaxial side of tepals with simple unicellular hairs and papillae. Septal nectaries external, present along upper abaxial side of ovary. Disc absent.

Androecium Stamens 3+3, inner stamens somewhat shorter than outer stamens. Filaments free from each other, adnate to mouth of perianth tube. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, perforate or microreticulate, granulate.

Gynoecium Pistil composed of three more or less connate carpels. Ovary inferior to slightly semi-inferior, trilocular, with a central canal. Style single, simple, filiform, long. Stigma punctate, type? Pistillodium absent.

Ovules Placentation axile to subbasal. Ovules two per carpel, anatropous, apotropous, bitegmic, crassinucellar to pseudocrassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument five to seven cell layers thick. Inner integument two cell layers thick. Obturator present (funicular?). Parietal cell formed from archesporial cell. Parietal tissue approx. three cell layers thick. Hypostase present. Nucellar cap two or three cell layers thick. Chalazal part of megasporangium relatively well developed. Megagametophyte monosporous, Polygonum type; with chalazal constriction. Antipodal cells ephemeral. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A one-seeded loculicidal capsule.

Seeds Aril absent. Seed coat exotestal. Exotesta with phytomelan layers on epidermal cell walls, palisade, with thin cell walls, remaining cells rounded. Mesotesta? Endotesta? Tegmen developed at micropyle, persistent. Perisperm not developed. Endosperm copious, ab initio with starch. Embryo?, chlorophyll? Cotyledon one? Cotyledon hyperphyll? Hypocotyl internode? Coleoptile? Germination?

Cytology n = 18

DNA

Phytochemistry Very insufficiently known. Lanaroflavone (a biflavone) and chelidonic acid present.

Use Unknown.

Systematics Lanaria (1; L. lanata; mountains in the Hottentots Holland in Western Cape to near Port Elizabeth in Eastern Cape).

Lanaria is sister-group to Hypoxidaceae.

The sieve tube plastids are of a unique type.

LAXMANNIACEAE Bubani

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Bubani, Fl. Pyren. 4: 111. 15-28 Feb 1902

Xerotaceae (Endl.) Hassk., Cat. Hort. Bot. Bogor.: 27. Oct 1844 [’Xerotideae’], nom. illeg.; Lomandraceae Lotsy, Vortr. Bot. Stammesgesch. 3: 761. 3-20 Sep 1911; Eustrephaceae Chupov in Bot. Žurn. 79(3): 7. Mar-Sep 1994

Genera/species 13–14/c 180

Distribution Tropical Asia, New Guinea, Australia, Tasmania, New Caledonia, New Zealand, South America, with their largest diversity in Australia.

Fossils Uncertain. Fossil pollen grains assigned to Cordyline are known from the Early Miocene onwards. Fossil leaves from the Eocene of South Australia (Paracordyline) are very similar to Australasiatic species of Cordyline, and leaf fossils from the Oligocene of Kerguelen resemble extant species of the genus from New Zealand.

Habit Bisexual or dioecious, usually perennial herbs (in Cordyline often fruticose or arborescent). Some species are xerophytic.

Vegetative anatomy Roots fibrous or fleshy (sometimes tuberous or stilt roots), in some species with a sclerenchymatous cortical zone outside root endodermis. Phellogen? Secondary lateral growth usually absent (anomalous secondary lateral growth present in stems of some species of Cordyline and Lomandra). Vessels usually present only in roots. Vessel elements with scalariform or simple perforation plates (in Acanthocarpus and Arthropodium in leaves also, then with scalariform perforation plates; in Eustrephus also in stem and leaves, with scalariform perforation plates); lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals, or P2cf type, with cuneate protein crystals and peripheral protein filaments (in Acanthocarpus P2ccl type, with cuneate and several additional loosely packed protein crystals). Nodes? Tanniniferous cells probably absent. Silica bodies absent. Idioblasts (raphide cells) with calciumoxalate raphides.

Trichomes Hairs unicellular or multicellular, uniseriate?, or absent.

Leaves Alternate (spiral or distichous), simple, entire, usually linear, sometimes terete or triangular in cross-section (in Cordyline sometimes differentiated into pseudopetiole and pseudolamina), with supervolute, conduplicate or flat to curved ptyxis. Stipules absent; leaf sheath well developed (in Laxmannia and Sowerbaea with ligule at sheath apex). Venation parallelodromous. Stomata usually anomocytic (sometimes paracytic or tetracytic). Cuticular wax crystalloids? Mesophyll with calciumoxalate raphides. Epidermal cells in Lomandra with calciumoxalate as styloid-like crystals (rhomboidal or polyhedral crystals present in leaf cells in some species of Lomandra and Romnalda). Leaf margin entire or spinulate-dentate.

Inflorescence Terminal, panicle, raceme-, umbel- or spike-like, sometimes with cymose partial inflorescences.

Flowers Actinomorphic, often small. Pedicel usually articulated, with pericladium. Hypogyny. Tepals 3+3, petaloid (in Thysanotus with inner tepals fimbriate), dry or fleshy, marcescent, usually free (sometimes connate at base). Septal nectaries infralocular (absent in some species). Disc absent.

Androecium Stamens 3+3, or three inner stamens (three outer stamens sometimes staminodial or absent). Filaments free from each other and usually from tepals (all or only inner stamens sometimes adnate to tepals; filaments in Arthropodium hairy). Anthers basifixed, centrifixed or dorsifixed, versatile?, tetrasporangiate, introrse or extrorse, usually longicidal (dehiscing by longitudinal slits; in Eustrephus and some species of Arthropodium and Thysanotus poricidal, dehiscing with apical pores). Tapetum usually secretory (in Eustrephus possibly amoeboid-periplasmodial), with binucleate cells. Staminodia sometimes three (in Arthropodium and Trichopetalum with staminodial appendage).

Pollen grains Microsporogenesis successive. Pollen grains usually monosulcate (in Acanthocarpus and Chamaexeros zonosulcate, with a single sulcus surrounding entire pollen grain and dividing it into two halves; in Lomandra sometimes spiraperturate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, in Acanthocarpus negatively reticulate, in Lomandra spinulate, echinate or negatively reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple. Stigmas one to three, Dry or Wet type. Pistillodium?

Ovules Placentation axile. Ovules one, several or numerous per carpel, anatropous to campylotropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal cell formed from archesporial cell (at least in Lomandra). Megasporangium with enlarged multilayered basal zone (proximal zone) with central ’Zuleitungsbahn’ consisting of axially elongated cells. Dermal chalazal cells enlarged. Nucellar cap present or absent. Megagametophyte monosporous, Polygonum typ. Nuclei of egg cell, synergids and antipodal cells enlarged. Synergids in some species with a filiform apparatus. Antipodal cells enlarged. Lateral megagametophyte haustoria present in at least Arthropodium. Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit Usually a loculicidal capsule (in Lomandra spicata a drupe; in Eustrephus a berry-like capsule) with persistent tepals.

Seeds Aril (formed from hilum) present in Murchisonia and Thysanotus. Strophiole (juicy, sweet) present in Eustrephus. Testa thin. Exotesta with or without phytomelan layer on epidermal cell walls. Mesotesta and endotesta often compressed or collapsed. Tegmen collapsed. Perisperm not developed. Endosperm copious, with lipids, aleurone and hemicellulose (starch absent); endosperm in Lomandra group and Eustrephus with thick, pitted cell walls. Embryo usually small, straight, curved or spirally twisted, chlorophyll? Cotyledon one, with apex sometimes transformed into haustorium, usually not photosynthesizing. Cotyledon hyperphyll? Hypocotyl internode in Cordyline long. Mesocotyl absent. Coleoptile usually absent (present in Sowerbaea). Radicula in Cordyline persistent, branched. Germination cryptocotylar?

Cytology n = 4 (Laxmannia, Sowerbaea); n = 6 (Chamaescilla); n = 7 (Chamaexeros, Lomandra); n = 8 (Acanthocarpus, Laxmannia, Lomandra); n = 10 (Eustrephus); n = 11 (Arthropodium, Chamaescilla, Murchisonia, Thysanotus); n = 19 (Cordyline) – Polyploidy frequently occurring. Chromosomes in Laxmannia, Sowerbaea and the Lomandra Group 2–7 µm long, or in Arthropodium and the Cordyline Group 0.5–2 µm long. Karyotype in Cordyline bimodal.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Steroidal saponins, naphthoquinones (in Lomandra), polyacetate derived arthroquinones (in Lomandra), phytosterols (in Cordyline), and chelidonic acid present. Polysaccharides in some genera stored as fructans. Seeds in Cordyline rich in linolic and oil acid based lipids. Flavonols, ellagic acid and alkaloids not found.

Use Ornamental plants, textile plants (Cordyline, Lomandra).

Systematics Laxmanniaceae are sister to [Asparagaceae+Ruscaceae].

The leaves in Xanthorrhoeaceae, some Lomandreae and in Dasypogonaceae are superficially similar. The leaves have sclerenchyma ribs extending from the inner envelopes of the vascular bundles. However, in Xanthorrhoea they arise from the mesophyll, whereas Dasypogonaceae lack such an envelope.

The subdivision below is according to Stevens (2001 onwards). A detailed phylogenetic analysis is in demand.

Lomandreae Engl. in Engler et Prantl, Nat. Pflanzenfam. II, 5: 18. 26 Mar 1887

4–5/c 65. Lomandra (c 50; New Guinea, Australia, New Caledonia), Romnalda (3; New Guinea, Queensland), Chamaexeros (4; southwestern Western Australia), Acanthocarpus (7; coastal regions in Western Australia); Xerolirion (1; X. divaricata; southwestern Western Australia)? – New Guinea, Australia, New Caledonia. Tubers absent. Leaves distichous, flat or curved. Sclerenchyma ribs of leaf extending from inner envelope of vascular bundle to surface; outer envelope with enlarged cells. Leaf margin sometimes serrate. Pedicel usually articulated. Pollen grains in Lomandra sometimes spiraperturate. Stigma Wet type. Ovules one or two per carpel. Nucellar cap present. Megasporangium with axially arranged central vascular passage. Tegmen brown, collapsed. Phytomelan absent. Endosperm with hemicellulose. n = 7, 8. Chromosomes 2–7 μm long.

Xerolirion divaricata has solitary terminal female flowers, male flowers in cymes, non-articulated pedicels, one ovule per carpel, lacks silica bodies, and has cell walls containing ferulates. Its systematic position is obscur.

Laxmannioideae Thorne et Reveal in Bot. Rev. 73(2): 82. 29 Jun 2007 [‘Laxmannieae’ Engl. 1887]

8/c 95. Arthropodium (16; Madagascar, Australia, New Caledonia, New Zealand), Murchisonia (2; southern and western Australia), Thysanotus (c 50; southern China to New Guinea, Australia), Trichopetalum (1–2; T. plumosum; Chile), Eustrephus (1; E. latifolius; southern and eastern New Guinea, eastern Queensland, eastern New South Wales, eastern Victoria, New Caledonia), Laxmannia (13; Australia), Sowerbaea (5; southwestern and eastern Australia), Chamaescilla (4; southwestern Western Australia, southern South Australia, Victoria, Tasmania). – Southeast Asia to Australia, New Caledonia, New Zealand, Chile. Vesicular-arbuscular mycorrhiza present. Storage roots present. Mucilage cells present. Leaves spiral, with supervolute or conduplicate ptyxis. Leaf sheath in Laxmannia and Sowerbaea with apical ligule. Flowers fasciculate or solitary. Pedicel often articulated. Anthers sometimes poricidal. Stigma Wet type. Megasporangium with axially oriented vascular tissue. Aril present in Murchisonia and Thysanotus. Exotesta often papillate, remaining testal layers cellular. Phytomelan present. Tegmen thin. Endosperm thin-walled. n = 4, 11. Chromosomes 0.5–2 μm long.

Cordylineae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 227. 20 Jul 1943

1/c 20. Cordyline (c 20; the Mascarene Islands, India eastwards to New Guinea, tropical Australia, New Zealand, islands in the Pacific; one species in tropical America from southern Brazil to Bolivia and northern Argentina). – Sometimes arborescent. Storage roots present. Mucilage cells present. Leaves spiral, in some species differentiated into pseudopetiole and pseudolamina, with supervolute or conduplicate ptyxis. Vascular bundles complex, amphivasal. Stomata paracytic; subsidiary cells with oblique divisions. Leaf sheath sometimes with apical ligule. Flowers solitary. Pedicel not articulated. Testa with phytomelan layer. n = 3, 6, 19. Chromosomes 0.5–2.4 μm long. Karyotype bimodal.

ORCHIDACEAE Juss.

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de Jussieu, Gen. Plant.: 64. 4 Aug 1789 [’Orchideae’], nom. cons.

Orchidales Raf., Anal. Nat.: 197. Apr-Jul 1815 [’Orchidia’]; Orchidopsida Bartl., Ord. Nat. Plant.: 24, 54. Sep 1830 [’Orchideae’]; Apostasiaceae Lindl., Nix. Plant.: 22. 17 Sep 1833 [’Apostasieae’], nom. cons.; Cypripediaceae Lindl., Nix. Plant.: 22. 17 Sep 1833 [’Cypripedieae’]; Neottiaceae Horan., Prim. Lin. Syst. Nat.: 50. 2 Nov 1834 [’Neottiaceae (Orchideae)’]; Apostasiales Blume in C. F. P. von Martius, Consp. Regn. Veg.: 10. Sep-Oct 1835 [’Apostasieae’]; Vanillaceae Lindl., Key Bot.: 73. 15-30 Sep 1835; Limodoraceae Horan., Char. Ess. Fam.: 44. 30 Jun 1847 [’Limodoraceae s. Orchideae’]; Cohniaceae Pfeiffer, Nomencl. Bot. 1(2): 815. ante 24 Jan 1873, nom. illeg.; Liparidaceae Vines, Stud. Text-book Bot. 2: 567. Mar 1895 [’Liparidinae’]; Ophrydaceae Vines, Stud. Text-book. Bot. 2: 566. Mar 1895 [‘Ophrydinae’]; Orchididae Heintze, Cormofyt. Fylog.: 10. 1927 [‘Orchidiformes’]; Neuwiediaceae (Burns-Bal. et V. A. Funk) R. Dahlgren ex Reveal et Hoogland in Bull. Mus. Natl. Hist. Nat., sér. IV, sect. B, Adansonia, 13: 91. 4 Oct 1991; Orchidanae Doweld, Tent. Syst. Plant. Vasc.: lviii. 23 Dec 2001

Genera/species 785–795/22.000–23.000

Distribution Cosmopolitan except polar regions, with the highest diversity in tropical and subtropical Asia and tropical Central and South America.

Fossils Meliorchis caribea is represented by a pollinium attached to a bee preserved in Oligocene to Miocene amber from the Dominican Republic.

Habit Usually bisexual (rarely monoecious or dioecious), perennial herbs. Many species have rhizome, root tubers or tuberous stem. Numerous species (c. 70%) are epiphytes, often with internodes modified into swollen water-storing pseudobulbs, often with contractile aerial roots covered with a thick layer of dead water-absorbing tissue, velamen, formed from epidermis; other species are climbing. Some genera consist of achlorophyllous holoendoparasites on fungi (intracellular modified ectomycorrhiza with mostly basidiomycetes). Several genera (especially within Vandeae) lack photosynthesizing leaves, which are replaced by photosynthesizing roots.

Vegetative anatomy CAM physiology often present (especially in Epidendroideae). Roots fibrous or tuberous, with mycorrhiza and usually with water-absorbing velamen (especially in epiphytic representatives). Plants as young (protocorms; sometimes also as mature) mycotrophic parasites on basidiomycetes (Russulaceae, Ceratobasidiaceae, Sebacinaceae, Thelephoraceae, Tulasnellaceae, sometimes Atractiellomycetes; sometimes ascomycetes, i.a. Tuber), with modified ectomycorrhiza as hyphal coils, pelotones, in cortical cells; also arbuscular mycorrhiza with Glomales. Phellogen absent. Primary vascular tissue as scattered bundles, centrifugal. Secondary lateral growth absent. Vessels present in roots (sometimes also in stem, rarely in leaves). Vessel elements usually with simple perforation plates (sometimes scalariform; in leaves only scalariform); lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate prostein crystals. Nodes? Envelopes of vascular bundles with idioblasts, stegmata, containing silica bodies; envelopes with fibres. Mucilage cells usually present. Calciumoxalate as raphides.

Trichomes Absent or multicellular, uniseriate, eglandular or glandular.

Leaves Alternate (spiral or distichous), simple, entire, often linear (sometimes equitant), often succulent, sometimes scale-like, with convolute (supervolute), conduplicate or (conduplicate-)plicate ptyxis. Stipules absent; leaf sheath usually closed. Colleters sometimes present. Venation parallelodromous. Fibre bundles sometimes present. Stomata usually paracytic (sometimes anomocytic or tetracytic). Cuticular wax crystalloids usually absent (rarely as non-orientated rodlets or scales). Epidermis often with stegmata containing silica bodies. Mesophyll often with mucilaginous idioblasts containing calciumoxalate raphides. Leaf margin usually entire. Extrafloral nectaries sometimes present.

Inflorescence Terminal or axillary, spike, raceme, or panicle (sometimes umbel or head; flowers sometimes solitary). Extrafloral nectaries sometimes present on bracts or abaxial side of outer tepals.

Flowers Usually zygomorphic (in Apostasioideae almost actinomorphic; in Neuwiedia resupinate; in Apostasia non-resupinate). Flower usually resupinate (usually by 180°; in some species re-resupinate, in other species non-resupinate; female flowers in Catasetum non-resupinate, male flowers resupinate). Epigyny. Tepals 3+3, usually with imbricate aestivation, caducous, usually free (sometimes connate at base); outer tepals sepaloid or petaloid; upper (median) tepal in outer whorl abaxial (usually seemingly adaxial due to resupination); inner tepals usually petaloid; upper (median) tepal in inner whorl usually modified into adaxial labellum (usually seemingly abaxial due to resupination; in Cypripedioideae slipper-shaped; in Orchidoideae and Epidendroideae often spurred), often larger and formed in different way than remaining tepals and often with spur with or without nectar. Tepal nectaries usually present. Septal nectaries absent. Oil secretion sometimes present (sometimes with elaiophores). Osmophores often present. Disc absent.

Androecium Stamens one, two or three; one median in outer whorl and two laterals in inner whorl; in Apostasioideae median outer stamen fertile or staminodial and lateral inner stamens fertile; in Cypripedioideae median outer stamen staminodial and lateral inner stamens fertile; in Vanilloideae, Orchidoideae and Epidendroideae median outer stamen fertile and lateral inner stamens staminodial. When filaments two or three then connate at base, free from tepals and partially adnate to style; when filament one then entirely or partially (at least at base) and congenitally adnate to style into a gynostemium and sometimes adnate to tepals (epitepalous). Anthers basifixed or dorsifixed, non-versatile, tetrasporangiate, latrorse or introrse, longicidal (dehiscing by longitudinal slits; sometimes with appendage); thecae usually transformed into pollinia. Placentoid often present. Tapetum secretory, with usually uninucleate or binucleate (rarely multinucleate) cells. Staminodia two intrastaminal or one extrastaminal or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually inaperturate (sometimes monosulcate, disulcate, monoporate, diporate, triporate, tetraporate, ulcerate, sulculate, or foraminate), usually shed as tetrads and coherent into massulae or in two (to eight) pollinia with single pollen grains or tetrads adhered to each other through strands of sterile sporogenous material (in some genera shed as monads), bicellular at dispersal. Exine tectate or semitectate, with usually columellate (sometimes granular) infratectum, reticulate to perforate, rugulate, scabrate, verrucate, psilate or smooth (exine rarely absent).

Gynoecium Pistil composed of three connate carpels; median carpel abaxial. Ovary inferior, usually unilocular (in Apostasioideae and some Cypripedioideae trilocular). Style single, simple, solid or with a stylar canal. Stigma capitate, usually trilobate (rarely bilobate), papillate, Wet type; median stigmatic lobe usually transformed into a sterile rostellum with viscidium. Pistillodium absent.

Ovules Placentation usually parietal-marginal (when ovary unilocular; in Apostasioideae and some Cypripedioideae axile); placentae branched. Ovules usually c. 30 to more than 1.500 per carpel (in one species almost four million per ovary), anatropous, usually bitegmic (rarely unitegmic), usually tenuinucellar (sometimes crassinucellar). Funicle non-vascularized. Micropyle endostomal. Outer integument one or two cell layers thick. Inner integument one or two cell layers thick. Parietal cell not formed (parietal tissue absent). Hypostase present. Nucellar cap absent. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type, or tetrasporous, ? type), with development usually delayed until after pollination. Antipodal cells persistent. Endosperm development ab initio nuclear; endosperm usually not developing, or development very early arrested. Endosperm haustoria absent. Embryogenesis asterad or onagrad (sometimes possibly piperad). Polyembryony occurs in some genera.

Fruit Usually a loculicidal or septicidal capsule (sometimes indehiscent; rarely a berry).

Seeds Aril absent. Seeds usually extremely small (often less than 0,1 µm). Seed coat membranous, mainly exotestal. Testa hyaline. Phytomelan absent. Tegmen sometimes thick. Perisperm not developed. Endosperm little developed or absent. Embryo minute, usually undifferentiated (devoid of organs), usually with chlorophyll. Suspensor haustoria usually well developed, with various structures (often branched). Cotyledon one, rarely developed. Coleoptile absent. Germination taking place together with fungal hyphae; embryo developing into a tuberous protocorm with basal rhizoids. Radicula usually absent (rarely present and then ephemeral).

Cytology n = 9, 10, 12, 14–16, 18, 19, 24, 29 or more

DNA

Phytochemistry Flavonols, flavone-C-glycosides, flavonoid sulphates, pyrrolizidine alkaloids in the form of esters of arylic and aralkylic acids (above all in Kerosphaereae) and other alkaloids, and 6-hydroxyapigenin methyl ethers, and aliphatic monocarboxylic esters present. Tricine rare. Mannans present as storage carbohydrates. Chelidonic acid? Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Aluminium accumulation reported from Spathoglottis.

Use Ornamental plants, flavours and perfumes (Vanilla), medicinal plants.

Systematics Orchidaceae are sister-group to all other Iridales.

Apostasioideae are sister to the clade [Vanilloideae+[Cypripedioideae+[Orchidoideae+Epidendroideae]]].

The taxonomy below is according to Chase & al. (2015).

Apostasioideae Horan., Char. Ess. Fam.: 46. 17Jun 1847 [‘Apostasieae‘]

2/14. Neuwiedia (8; Yunnan to Malesia), Apostasia (6; Sri Lanka, northeastern India, Japan, Southeast Asia, Malesia, northeastern Queensland, Melanesia). – Sri Lanka, northeastern India, Japan, Southeast Asia, Malesia, northeastern Queensland, Melanesia. Vessels present in roots and sometimes in stem. Vessel elements with simple or scalariform perforation plates. Leaves spiral, with plicate ptyxis. Stomata tetracytic. Flowers almost actinomorphic; in Neuwiedia resupinate; in Apostasia non-resupinate, without labellum. Simultaneous initiation of inner tepals. Tepals apiculate, carinate. Stamens in Neuwiedia three fertile (staminodium absent); in Apostasia two lateral inner fertile stamens and one median outer staminodium. Pollen grains sometimes operculate. Ovary trilocular. Style entire. Placentation axile. Micropyle bistomal. Megagametophyte in Neuwiedia disporous, Allium type. Seed coat endotestal. Exotegmen in Neuwiedia sclerified. Outer periclinal cell walls collapsing. n = 24. Chromosomes small. Plastid gene matK possibly transition between functional gene and pseudogene.

[Vanilloideae+[Cypripedioideae+[Orchidoideae+Epidendroideae]]]

Velamen often present. Tilosomes (‘fibrous bodies‘, ‘rod bodies‘; lignified appendages from cell walls of innermost cell layer of root velamen adjacent to exodermis) often present. Vessel elements usually with scalariform perforation plates. Leaves spiral or distichous. Flowers strongly zygomorphic, with well developed labellum. Tepal nectaries sometimes present. Style and filaments almost entirely congenitally fused into gynostemium. Pollen grains viscid. Stigma asymmetric. Placentation parietal. Ovules not entirely developed at pollination (fertilization often delayed up to several months). Seeds dispersed prior to embryo maturation. Radicula absent or ephemeral.

Vanilloideae (Lindl.) Szlach. in Fragm. Forist. Geobot., Suppl. 3: 48. 1995

14/c 250. Pogonieae Pfetzer ex Garay et dunsterv., Venez. Orchids Ill. 2: 28. 1961. Duckeella (3; tropical South America), Cleistes (64; tropical America to Florida), Cleistesiopsis (2; eastern and southeastern United States), Isotria (2; eastern United States), Pogonia (5; temperate Asia, North America). – Vanilleae Blume in Rumphia 1: 196. Dec 1837. Epistephium (21; northern South America), Lecanorchis (c 20; tropical Asia, Japan), Clematepistephium (1; C. smilacifolium; New Caledonia), Eriaxis (1; E. rigida; New Caledonia); Vanilla (106; tropical and subtropical regions on both hemispheres), Erythrorchis (2; tropical Asia, tropical Australia), Pseudovanilla (≥8; Malesia, tropical Australia, Pacific islands), Cyrtosia (5; tropical Asia, Japan), Galeola (6; tropical Asia). – Pantropical, few species in eastern North America, with their largest diversity in tropical Asia. CAM physiology sometimes present. Silica bodies absent. Stomata usually tetracytic. Simultaneous initiation of inner tepals? Tepals often carinate, sometimes persistent. Median outer stamen fertile and lateral inner stamens staminodial. Anther incumbent due to strong extension of apical connective. Pseudopollinia soft (pollinia absent). Pollen grains sometimes polyporate, often shed as tetrads. Viscidium absent. Rostellum (part of median stigmatic lobe) present. Placentation usually parietal (sometimes axile). Seeds fusiform, crustose. Seed coat exotestal, with outer parietal wall well developed. Tegmen persistent. Endosperm up to 16-nucleate. n = 9, 10, 12, 14–16, 18 and more. Chromosomes comparatively large. Plastid gene matK pseudogene. – Vanilloideae are sister-group to the remaining Orchidaceae.

[Cypripedioideae+[Orchidoideae+Epidendroideae]]

Cypripedioideae Lindl. ex Endl., Gen. Plant.: 220. Jun 1837 [’Cypripedieae’]

5/c 170. Cypripedium (51; temperate regions on the Northern Hemisphere), Paphiopedilum (86; tropical Asia to southern China, New Guinea and southwestern Pacific islands), Selenipedium (5; tropical America), Phragmipedium (26; tropical America), Mexipedium (1; M. xerophyticum; Mexico). – Tropical, subtropical and temperate regions on the Northern Hemisphere to East Malesia, southern India, southwestern Pacific and southwards to central South America. Leaves with conduplicate or plicate ptyxis. Stomata often paracytic. Simultaneous initiation of inner tepals. Tepals sometimes caducous. Two abaxial outer tepals connate. Labellum strongly saccate. Two lateral inner stamens fertile. Median outer stamen staminodial. Tapetum with binucleate cells. Microsporogenesis successive? Pollen grains operculate, sometimes shed as tetrads. Ovary sometimes trilocular. Median stigmatic lobe larger than lateral lobes. Placentation usually parietal (sometimes axile). Micropyle sometimes bistomal. Megagametophyte sometimes disporous, octocellular, Allium type. n = 9 or more. Chromosomes comparatively large. Plastid gene matK pseudogene. – Cypripedioideae are sometimes sister-group to Vanilloideae or to the clade [Vanilloideae+[Orchidoideae+Epidendroideae]]. Selenipedium, in particular, have a number of plesiomorphies, e.g. pollen grains shed as monads, trilocular ovary, fleshy fruit, and hard dark testa, suggesting a position of Cypripedioideae at least as basal as Vanilloideae. Yet, I have followed the most commonly recovered topology here.

[Orchidoideae+Epidendroideae]

Floral primordium transversely elliptic-oval. Tepals not caducous. Median outer stamen fertile. Two lateral inner stamens usually absent (sometimes staminodial). Pollinia adnate to viscidium (part of median stigmatic lobe). Pollinarian stipe formed from caudicula (part of anther), tegula (epidermis on rostellum), or hamulus (apex of rostellum). Microsporogenesis simultaneous, with tetraedric tetrads. Pollen grains porate or ulcerate, usually shed in tetrads. Median carpel developed prior to lateral carpels. Rostellum (part of median stigmatic lobe) present. Endosperm not developing. Tegmen not persistent. n = 9 or more (often 19).

Orchidoideae Eaton, Bot. Dict., ed. 4: 29. Apr-Mai 1836 [‘Orchideae’]

c 210/c 3.630. Codonorchideae P. J. Cribb in P. J. Cribb et P. J. Kores, Lindleyana 15: 169. 29 Sep 2000. Codonorchis (2; Brazil to southern South America, the Falkland Islands). – Orchideae Small, Man. S.E. Fl.: 363. 30 Nov 1933. Brownleeinae H. P. Linder et H. Kurzweil in Ann. Missouri Bot. Gard. 81: 710. 1994. Brownleea (8; tropical and southern Africa, Madagascar), Disperis (78; Africa, Madagascar, the Mascarene Islands, India, Sri Lanka, Thailand, China, Japan, the Ryukyu Islands, Taiwan, the Philippines, Java, the Lesser Sunda Islands, New Guinea, the Caroline Islands). – Coryciinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [‘Corycieae’].Ceratandra (6; Western and Eastern Cape), ‘Corycium’ (15; southern Africa and northwards to Malawi and southern Tanzania; non-monophyletic), Evotella (1; E. rubiginosa; Western Cape), ‘Pterygodium’ (19; southern Africa, one species in Tanzania, with their highest diversity in Western Cape and the Drakensberg in KwaZulu-Natal; non-monophyletic). – Pachites clade Pachites (2; Western Cape). – Disinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [‘Diseae’]. Disa (c 180; Africa, Madagascar, the Mascarene Islands, the Arabian Peninsula), Huttonaea (5; Eastern Cape, KwaZulu-Natal, Mpumalanga, Free State).– Orchidinae Reveal in Phytoneuron 2012-33: 2. 9 Apr 2012. Satyrium (c 85; tropical and southern Africa, Madagascar, the Arabian Peninsula, India, Sri Lanka, Burma, China), Peristylus (c 100; China to islands in the Pacific), Pecteilis (8; tropical and East Asia; in Habenaria?), Hsenhsua (1; H. chrysea; easter Himalayas, Bhutan, Yunnan, Xizang), Herminium (c 20; Europe, temperate Asia), Androcorys (10; Himalayas, China, Honshu in Japan), Porolabium (1; P. biporosum; China), Gennaria (1; G. diphylla; Macaronesia, western Mediterranean; in Habenaria?), Nujiangia (1; N. griffithii; Himalayas),‘Habenaria’ (c 835; tropical and subtropical regions on both hemispheres; paraphyletic), Bonatea (13; eastern Africa southwards to Western Cape, the Arabian Peninsula; in Habenaria?), Sirindhornia (3; Burma, Yunnan, Thailand), Tsaiorchis (1; T. neottianthoides; southeastern Yunnan, Guangxi), Hemipilia (13; Himalayas, southeastern Tibet, East Asia, Thailand),‘Ponerorchis’ (c 55; Himalayas, China, Japan; paraphyletic), Ophrys (c 35; Europe, the Mediterranean, North Africa, western Asia), Serapias (13; the Azores, the Mediterranean), Orchis (21; temperate regions on the Northern Hemisphere and southwards to southwestern China and northern India), Gymnadenia (23; Europe, temperate Asia, eastern North America), Coeloglossum (1; C. viride; temperate regions on the Northern Hemisphere), Dactylorhiza(c 40; Europe, Macaronesia, the Mediterranean, temperate Asia, Alaska), Pseudorchis(2; Europe, eastern North America), Galearis (10; Himalayas, Tibet, Burma, China, the Korean Peninsula, Japan, Russian Far East, Kamtchatka, temperate North America, western Greenland),’Platanthera’ (c 135; temperate to tropical regions on the Northern Hemisphere southwards to mountains in Malesia; polyphyletic), Anacamptis (11; Europe, the Mediterranean and eastwards to Iran), Bartholina (2; southern Namibia, Northern, Western and Eastern Cape), Benthamia (c 30; Madagascar, the Mascarene Islands?), Bhutanthera (5; Nepal, eastern Himalayas to southwestern China), Brachycorythis (c 35; tropical and southern Africa, tropical Asia), Centrostigma (3; southern tropical and southern Africa), Chamorchis (1; C. alpina; northern Scandinavia, the Carpathians, the Alps), Cynorkis (c 155; eastern Africa, Madagascar, the Mascarene Islands, with their highest diversity in Madagascar), Diplomeris (3; Himalayas), Dracomonticola (1; D. virginea; the Drakensberg in KwaZulu-Natal and Lesotho), Frigidorchis (1; F. humidicola; Qinghai in southern China), Himantoglossum (11; Europe, the Canary Islands, the Mediterranean and eastwards to western Iran, North Africa), Holothrix (c 45; tropical and southern Africa, the Arabian Peninsula), Megalorchis (1; M. regalis; eastern Madagascar), Neobolusia (3; central, eastern and southern Africa), Neotinea (4; Europe, Macaronesia, the Mediterranean), Odisha (1; O. cleistantha; Orissa in India; in Habenaria?), Oligophyton (1; O. drummondii; Zimbabwe), Physoceras (12; Madagascar), Platycoryne (c 20; tropical Africa, Madagascar), Roeperocharis (5; tropical East Africa), Schizochilus (11; eastern and southeastern Africa), Silvorchis (3; Vietnam, Java), Stenoglottis (7; eastern South Africa, Swaziland, one species in eastern Africa to Tanzania), Steveniella (1; S. satyroides; Crimea to western Asia), Thulinia (1; T. albolutea; Nguru Mountains in Tanzania), Traunsteinera (2; Europe, the Mediterranean), Tylostigma (8; Madagascar), Veyretella (2; Gabon). – Cranichideae Pfeiff., Nomencl. Bot. 1(2): 901. ante 3 Oct 1873 (polyphyletic). Chloraeinae Pfitzer, Entwurf. Anordn. Orch.: 98. Jan-Apr 1887 [’Chloraeeae’]. Bipinnula (11; Brazil, Uruguay, Argentina, southern Chile), ‘Chloraea‘ (52; southern Chile, southern Argentina), Gavilea (15; southern Chile, southern Argentina).– Pterostylidinae Pfitzer, Entwurf. Anordn. Orch.: 97. Jan-Apr 1887 [’Pterostylideae’]. Achlydosa (1; A. glandulosa; New Caledonia), Pterostylis (c 210; East Malesia, Queensland to South Australia, Tasmania, New Caledonia, New Zealand). – Goodyerinae Ridl., Mat. Flora Malay Penins. 1: 12. 1907. Aenhenrya (1; A. rotundifolia; southern India), Anoectochilus (43; tropical Asia to the Hawaiian Islands), Aspidogyne (c 60; tropical America), Chamaegastrodia (3; Himalayas, East Asia), Cheirostylis (53; tropical and subtropical regions of the Old World), Cystorchis (c 20; Thailand and eastwards to Micronesia), Danhatchia (1; D. australis; New Zealand), Dossinia (1; D. marmorata; Borneo), Erythrodes (26; tropical Asia, New Caledonia to Samoa and Tonga), Eurycentrum (7; New Guinea, the Solomon Islands, Vanuatu), Gonatostylis (2; New Caledonia), Goodyera (c 100; temperate regions on the Northern Hemisphere, Mozambique, Madagascar, tropical Asia), Halleorchis (1; H. aspidogynoides; Cameroun, Gabon), Herpysma (1; H. longicaulis; southern and southeastern Asia to Sumatra), Hetaeria (c 30; Sri Lanka, Southeast Asia, Malesia and eastwards to Tahiti), Hylophila (7; Thailand and eastwards to the Solomon Islands), Kreodanthus (14; tropical America), Kuhlhasseltia (9; the Korean Peninsula, Japan and southwards to New Guinea), Lepidogyne (1; L. longifolia; Malesia), Ludisia (1; L. discolor; Southeast Asia, West Malesia), Macodes (11; Japan, Vietnam, Malesia to Vanuatu), Microchilus (c 140; tropical America), Myrmechis (17; Japan, tropical Asia), Odontochilus (c 25; Himalayas, tropical Asia, the Hawaiian Islands), Orchipedum (3; West Malesia), Pachyplectron (3; New Caledonia), Papuaea (1; P. reticulata; New Guinea), Platylepis (17; tropical and southern Africa, Madagascar, the Mascarene Islands, the Seychelles, the Moluccas, Tahiti), Rhamphorhynchus (1; R. mendoncae; Brazil), Rhomboda (22; Himalayas, tropical Asia and eastwards to New Caledonia), Schuitemania (1; S. merrillii; the Philippines), Stephanothelys (5; the Andes), Vrydagzynea (43; tropical Asia, Taiwan, western Pacific islands), Zeuxine (c 75; tropical and subtropical regions in the Old World), Zeuxinella (1; Z. vietnamica; Vietnam). – Galeottiellinae Salazar et M. W. Chase in Lindleyana 17: 172. 9 Oct 2002. Galeottiella (6; Mexico, Guatemala). – Manniellinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 572, 585. 22 Jul 1926 [’Mannielleae’]. Manniella (2; tropical West Africa). – Spiranthinae Lindl. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 385, Comm. 288. 17-20 Aug 1842 [’Spiranthideae’].Cotylolabium (1; C. lutzii; Espírito Santo and Minas Gerais in southeastern Brazil);Aracamunia (1; A. liesneri; southern Venezuela), Aulosepalum (7; Mexico to Costa Rica), Beloglottis (7; southeastern United States and southwards to Brazil), Brachystele (21; tropical America), Buchtienia (3; tropical South America), Coccineorchis (7; Mexico, Central America, northwestern South America), Cotylolabium (1; C. lutzii; Brazil), Cybebus (1; C. grandis; Colombia), Cyclopogon (c 85; tropical and subtropical America), Degranvillea (1; D. dermatoptera; French Guiana), Deiregyne (18; Mexico, Guatemala), Dichromanthus (4; Texas, Mexico, Central America), Eltroplectris (13; tropical America, Florida), Eurystyles (20; tropical America), Funkiella (27; Texas, Mexico, Guatemala, Costa Rica), Hapalorchis (10; tropical and subtropical America), Helonoma (4; Guayana Highlands, the Andes in Peru), Kionophyton (4; Mexico, Guatemala), Lankesterella (11; tropical America), Lyroglossa (2; tropical America), Mesadenella (7; tropical America), Mesadenus (7; Florida, Central America, the West Indies, southeastern Brazil), Nothostele (2; Brazil), Odontorrhynchus (6; the Andes), Pelexia (77; tropical and subtropical America), Physogyne (3; Mexico), Pseudogoodyera (1; P. wrightii; Central America, Cuba), Pteroglossa (11; South America), Quechua (1; Q. glabrescens; Ecuador, Peru), Sacoila (7; tropical and subtropical America), Sarcoglottis (c 50; tropical America), Sauroglossum (11; South America), Schiedeella (c 25; Arizona and southwards to tropical South America), Skeptrostachys (13; tropical South America), Sotoa (1; S. confusa; western Texas, northern and central Mexico), Spiranthes (c 35; temperate regions on the Northern Hemisphere, Malesia, Australia, Pacific islands, tropical America), Stalkya (1; S. muscicola; the Andes in Venezuela), Stenorrhynchos (5; tropical America), Svenkoeltzia (3; Mexico), Thelyschista (1; T. ghillanyi; eastern Brazil). – Discyphinae Salazar et Van den Berg in Phytotaxa 173(2): §136. 25 Jun 2014. Discyphus (1; D. scopulariae; tropical America). – Cranichidinae Lindl. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 384, Comm. 288. 17-20 Aug 1842 [’Cranichideae’].Aa (c 25; the Andes), Altensteinia (8; the Andes), Baskervilla (6; the Andes, southern Brazil), Cranichis (53; Florida and southwards to the Andes), Fuertesiella (1; F. pterichoides; eastern Cuba, Dominican Republic), Galeoglossum (3; Mexico), Prescottia (26; tropical America), Gomphichis (24; Costa Rica, Brazil, the Andes), Myrosmodes (12; the Andes), Ponthieva (c 65; tropical and subtropical America), Porphyrostachys (2; the Andes in Ecuador and Peru), Pseudocentrum (7; tropical America), Pterichis (c 20; Costa Rica, Jamaica, the Andes), Solenocentrum (4; Costa Rica, Ecuador), Stenoptera (7; the Andes). – Diurideae Endl. ex Butzin in Willdenowia 6: 311. 22 Oct 1971. Rhizanthellinae R. S. Rogers in J. Roy. Soc. W. Australia 15: 4. 17 Oct 1928. Rhizanthella (2; southwestern Western Australia, eastern Australia). – Acianthinae Schltr. in Bot. Jahrb. Syst. 45: 381. 21 Feb 1922.Acianthus (20; New Guinea, eastern Queensland to southeastern South Australia, Tasmania, New Caledonia, New Zealand), Corybas (c 130; Japan, tropical Asia, eastern Queensland to southeastern South Australia, Tasmania, New Zealand, Macquarie Island, Polynesia), Cyrtostylis (5; southwestern Western Australia, southeastern Queensland to southeastern South Australia, Tasmania, New Zealand), Stigmatodactylus (10; the Himalayas, China, Japan, Indonesia, New Guinea, the Solomon Islands), Townsonia (2; Tasmania, New Zealand). – Caladeniinae Pfitzer, Entwurf. Anordn. Orch.: 97. Jan-Apr 1887 [’Caladenieae’]. Adenochilus (2; eastern New South Wales, New Zealand), Aporostylis (1; A. bifolia; New Zealand, Stewart Island, Chatham Islands, Auckland Islands, Campbell Island, the Antipode Islands),’Caladenia’ (c 270; Australia, one species, C. catenata, also in Malesia, New Caledonia, New Zealand, Antipode Island and Chatham Islands; paraphyletic), Cyanicula (10; southwestern Western Australia, southeastern Queensland, eastern New South Wales, Victoria; in Caladenia?), Elythranthera (2; southwestern Western Australia; in Caladenia?), Ericksonella (1; E. saccharata; southwestern Western Australia; in Caladenia?), Eriochilus (9; southwestern Western Australia, southeastern Australia, Tasmania), Glossodia (2; southeastern Queensland to southeastern South Australia, Tasmania; in Caladenia?), Leptoceras (1; L. menziesii; southwestern Western Australia, southeastern South Australia, Victoria, Tasmania; in Caladenia?), Pheladenia (1; P. deformis; southwestern Western Australia; in Caladenia?), Praecoxanthus (1; P. aphyllus; southwestern Western Australia). – Prasophyllinae Schltr. in Bot. Jahrb. Syst. 45: 381. 21 Feb 1911. Genoplesium (47; eastern Queensland to southern Western Australia, Tasmania, New Caledonia, New Zealand), Microtis (19; China, Japan to southwestern Western Australia, New Zealand), Prasophyllum (c 130; southeastern Queensland to southwestern Western Australia, Tasmania, New Zealand). – Diuridinae Lindl. ex Meisn., Plat. Vasc. Gen.: Tab. Diagn. 387, Comm. 289. 17-20 Aug 1842 [‘Diurideae‘]. Diuris (c 70; Timor, Australia), Orthoceras (2; southeastern Queensland to southeastern South Australia, Tasmania, New Caledonia, New Zealand). – Cryptostylidinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 570, 583. 22 Jul 1926 [‘Cryptostylideae‘]. Coilochilus (1; C. neocaledonicus; New Caledonia), Cryptostylis (c 25; Taiwan, tropical Asia and eastwards to Australia, New Caledonia, Vanuatu, New Zealand and Samoa). – Drakaeinae Schltr. in Bot. Jahrb. Syst. 45: 381. 21 Feb 1911. Arthrochilus (c 15; New Guinea, tropical Australia to northeastern New South Wales), Caleana (1; C. major; southwestern Western Australia, southeastern Queensland to South Australia, Tasmania), Chiloglottis (c 25; southeastern Queensland to southern Victoria, Tasmania, New Zealand), Drakaea (10; southwestern Western Australia), Paracaleana (13; eastern, southern and western Australia, Tasmania, North Island in New Zealand, with their highest diversity in southwestern Western Australia), Spiculaea (1; S. ciliata; southwestern Western Australia, southeastern New South Wales, southern Victoria). – Megastylidinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 572, 585. 22 Jul 1926 [‘Megastylideae‘].Burnettia (1; B. cuneata; southeastern New South Wales, southern Victoria, Tasmania), Leporella (1; L. fimbriata; southwestern Western Australia, southeastern South Australia, Victoria), Lyperanthus (2; southwestern Western Australia, southeastern Australia, Tasmania, New Zealand),’Megastylis’ (7; New Caledonia, Vanuatu; polyphyletic), Pyrorchis (2; southwestern Western Australia, southeastern South Australia, eastern New South Wales, Victoria, Tasmania), Rimacola (1; R. elliptica; eastern New South Wales), Waireia (1; W. stenopetala; New Zealand). – Thelymitrinae Lindl. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 387, Comm. 289. 17-20 Aug 1842 [’Thelymitrideae’]. Calochilus (27; New Guinea, southeastern Queensland to southeastern South Australia, southwestern Western Australia, Tasmania, New Caledonia, New Zealand), Epiblema (1; E. grandiflorum; southwestern Western Australia), Thelymitra (c 110; Malesia, eastern Queensland to southeastern South Australia, southwestern Western Australia, Tasmania, New Caledonia, New Zealand). – Cosmopolitan, with their highest diversity in temperate regions. Mycoheterotrophy absent. Stem or root tuber frequent. Stem and leaf sclerenchyma rare (in leaves as fibre bundles or associated with vascular bundles). Stegmata with silica bodies absent. Leaves spiral or distichous. Stomata usually anomocytic. Anther usually erect (sometimes incumbent). Pollinia present, soft, sectile. Staminodia reduced. Hamulus (pollinium stalk developed from modified rostellum apex) sometimes present. n = 12–24. Chromosomes small. Plastid gene matK pseudogene.

Epidendroideae Lindl. ex Endl., Gen. Plant.: 193. Jun 1837 [‘Epidendreae’]

555–560/c 18.000? Neottieae Lindl., Orchid. Select.: 7, 9. Jan 1826. Aphyllorchis (22; Sri Lanka to Japan and eastern Queensland), Cephalanthera (19; temperate regions on the Northern Hemisphere), Epipactis (c 50; Europe, tropical Africa, temperate Asia, Himalayas, Southeast Asia), Limodorum (3; the Mediterranean and eastwards to Iran), Neottia (c 65; Europe, temperate Asia, North America), Palmorchis (21; Central America, tropical South America). – Sobralieae Pfitzer, Entwurf. Anordn. Orch.: 99. Jan-Apr 1887 [’Sobraliinae’]. Elleanthus (c 110; tropical America; incl. Sertifera?), Sertifera (7; the Andes; in Elleanthus?), Sobralia (c 150; tropical America). – Tropidieae Dressler in Telopia 2: 422. 13 Oct 1983. Corymborkis (6; tropical Asia and eastwards to islands in the Pacific), Kalimantanorchis (1; K. nagamasui; western Borneo), Tropidia (c 30; tropical Asia to Taiwan and western Pacific islands, Florida, Central America, the West Indies). – Wullschlaegelieae Dressler in Telopea 2: 422. 13 Oct 1983.Wullschlaegelia (2; tropical America). – Triphoreae Dressler in Selbyana 5: 204. 27 Dec 1979. Diceratostele (1 D. gabonensis; western and central tropical Africa); Monophyllorchis (1; M. microstyloides; Colombia, Ecuador), Pogoniopsis (2; Brazil), Psilochilus (7–9; tropical America), Triphora (18; temperate to tropical America). – Xerorchideae P. J. Cribb in Kew Bull. 60: 143. Jul 2005. Xerorchis (2; northern South America). – Gastrodieae Lindl., Orchid. Select.: 7, 11. Jan 1826. Auxopus (4; tropical Africa), Didymoplexiella (7; Southeast Asia and eastwards to Japan and Borneo), Didymoplexiopsis (1; D. khiriwongensis; Thailand, Laos, Vietnam, Hainan; in Didymoplexiella?), Didymoplexis (17; tropical and southeastern Africa, Madagascar, Southeast Asia, Malesia), Gastrodia (c 60; East Asia, tropical Asia and eastwards to southwesternmost and southeastern Australia, Tasmania and New Zealand), Uleiorchis (2; tropical America). – Nervilieae Dressler in Lindleyana 5: 124. 29 Jun 1990.Nervilia (67; tropical and southern Africa, Madagscar, the Arabian Peninsula, tropical Asia eastwards to New Guinea and northern Australia); Epipogium (2–5; Europe, temperate to tropical Asia and eastwards to eastern Queensland and northeastern New South Wales), Stereosandra (1; S. javanica; Southeast Asia, West Malesia).– Thaieae X. H. Jin et X. G. Xiang in Taxon 61: 52. 21 Feb 2012.Thaia (1; T. saprophytica; Thailand). – Arethuseae Lindl., Orchid. Select.: 7, 10. Jan 1826. Arethusinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [‘Arethuseae’]. Arundina (1; A. graminifolia; Himalayas and eastwards to islands in the Pacific), Arethusa (1; A. bulbosa; eastern North America), Eleorchis (1; E. japonica; Japan); Anthogonium (1; A. gracile; eastern Himalayas to Southeast Asia), Calopogon (5; eastern North America, the West Indies). – Coelogyninae Benth. in J. Linn. Soc. London, Bot. 18: 287. 21 Feb 1881 [’Coelogyneae’].Aglossorhyncha (13; East Malesia, New Guinea), Bletilla (5; temperate East Asia), Bracisepalum (2; Sulawesi), Bulleyia (1; B. yunnanensis; eastern Himalaya, southwestern China), Chelonistele (13; West Malesia, with their highest diversity on Borneo), Coelogyne (c 200; tropical Asia, southern China, western Pacific islands), Dendrochilum (c 280; Southeast Asia, Malesia), Dickasonia (1; D. vernicosa; northeastern India, Burma), Dilochia (8; Malesia), Entomophobia (1; E. kinabaluensis; northern Borneo), Geesinkorchis (4; West Malesia), Glomera (c 130; Malesia and eastwards to western Pacific islands), Gynoglottis (1; G. cymbidioides; Sumatra), Ischnogyne (1; I. mandarinorum; China), Nabaluia (3; northern Borneo), Otochilus (5; eastern Himalayas to Southeast Asia), Panisea (7; northeastern India to Southeast Asia), Pholidota (31; India to southern China and Southeast Asia, Malesia to New Guinea, tropical and eastern Australia, Pacific islands), Pleione (16–22; Nepal to Taiwan and Thailand), Thunia (5; Himalayas to Burma). – Malaxideae Lindl., Collect. Bot. 8: 7. Jan 1826. Dendrobiinae G. Don in R. Sweet, Hort. Brit., ed. 3: 638. late 1839 [’Dendrobieae’]. Bulbophyllum (c 1.870; tropical and subtropical regions on both hemispheres), Chaseella (1; C. pseudohydra; Zimbabwe; in Bulbophyllum?)?, Dendrobium (c 1.510; tropical and subtropical regions in Asia, tropical and eastern Australia, Tasmania and the Pacific), Drymoda (3; Burma, Thailand, Laos, New Guinea; in Bulbophyllum?)?, Genyorchis (10; tropical West and Central Africa; in Dendrobium?)?, Jejosephia (1; J. pusilla; Assam)?, Monomeria (3; northern India, eastern Himalayas, Tibet, southern China, Southeast Asia; in Bulbophyllum?)?, Pedilochilus (c 35; New Guinea, the Solomon Islands, Vanuatu; in Dendrobium?)?, Saccoglossum (5; New Guinea; in Bulbophyllum?)?, Trias (14; northeastern India, the Andaman and Nicobar Islands, Burma, Thailand, Indochina, Borneo; in Bulbophyllum?)? – Malaxidinae Benth. et Hook. f., Gen. Plant. 3: 463, 465. 14 Apr 1883. Alatiliparis (5; Sumatra, Java), Crepidium (c 260; islands in the Indian Ocean, India, China, Southeast Asia, Malesia to New Guinea, northern Australia, islands in the Pacific), Crossoglossa (26; Central America, tropical South America), Crossoliparis (1; C. wendlandii; Central America), Dienia (6; tropical and subtropical Asia, Malesia, northern Australia), Hammarbya (1; H. paludosa; temperate and subarctic regions on the Northern Hemisphere), Hippeophyllum (c 10; Malesia), ’Liparis’ (c 425; nearly cosmopolitan; non-monophyletic),’Malaxis’ (c 180; nearly cosmopolitan; non-monophyletic), Oberonia (c 325; tropical and southern Africa, Madagascar, the Mascarene Islands, India and eastwards to New Guinea and northern Australia), Oberonioides (2; China, Thailand), Orestias (4; tropical Africa), Stichorkis (8; islands in the Indian Ocean, India, Southeast Asia, New Guinea, islands in the Pacific), Tamayorkis (4; tropical America). – Cymbidieae Pfitzer, Entwurf. Anordn. Orch.: 105. Jan-Apr 1887 [’Cymbidiinae’]. Cymbidiinae Benth. in J. Linn. Soc. London, Bot. 18: 287. 21 Feb 1881 [’Cymbidieae’]. Acriopsis (9; Southeast Asia and eastwards to the Solomon Islands), Cymbidium (c 70; northwestern India to Japan and Australia), Grammatophyllum (12; Malesia to Polynesia), Porphyroglottis (1; P. maxwelliae; Malaysia, Sumatra, Borneo), Thecopus (2; tropical Asia), Thecostele (1; T. alata; Burma to West Malesia). – Cyrtopodiinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [’Cyrtopodieae’]. Cyrtopodium (47; Florida, tropical America).– Eulophiinae Benth. in J. Linn. Soc. London, Bot. 18: 287. 21 Feb 1881 [’Eulophieae’]. Acrolophia (7; Western Cape to KwaZulu-Natal), Ansellia (1; A. africana; tropical and southern Africa), Claderia (2; Thailand to New Guinea), Cymbidiella (3; Madagascar), Dipodium (c 25; Malesia to New Caledonia), Eulophia (c 200; tropical and subtropical regions on both hemispheres), Eulophiella (5; Madagascar), Geodorum (12; tropical Asia to western Pacific islands), Grammangis (2; Madagascar), Graphorkis (4; tropical Africa, Madagascar, the Mascarene Islands), Imerinaea (1; I. madagascarica; northern Madagascar), Oeceoclades (31; tropical Africa, tropical America northwards to the West Indies), Paralophia (2; Madagascar). – Catasetinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 577, 588. 22 Jul 1926 [’Cataseteae’]. Catasetum (c 175; tropical America), Clowesia (7; Central America), Cyanaeorchis (3; Brazil), Cycnoches (c 35; tropical America), Dressleria (11; Central America), Galeandra (38; Central America, tropical South America), Grobya (5; Brazil), Mormodes (c 80; tropical America). – Oncidiinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [’Oncidieae’]. Ada (17; Central America, the Andes in Colombia to Bolivia)?, Aspasia (7; tropical America), Brachtia (8; northwestern South America)?, Brassia (c 65; Florida, tropical America), Caluera (3; northern South America), Capanemia (9; tropical South America), Caucaea (9; northwestern South America), Centroglossa (5; Brazil, Peru, Paraguay), Chytroglossa (3; southeastern Brazil), Cischweinfia (11; Central America, northwestern South America), Comparettia (c 80; tropical America), Cuitlauzina (7; Mexico, Central America), Cypholoron (2; Venezuela, Ecuador), Cyrtochiloides (3; tropical America), Cyrtochilum (c 140; Venezuela and Colombia to Peru), Dunstervillea (1; D. mirabilis; Venezuela, Ecuador, Brazil), Eloyella (10; Panamá, northern South America), Erycina (7; Mexico, Central America, Trinidad, tropical South America), Fernandezia (c 50; southern Mexico, Central America, tropical South America), Gomesa (c 120; South America), Grandiphyllum (7; Brazil, Argentina), Hintonella (1; H. mexicana; Mexico), Hofmeisterella (2; Ecuador), Ionopsis (6; Florida, Mexico, Central America, the West Indies, tropical South America), Leochilus (12; tropical America), Lockhartia (28; tropical America), Macradenia (11, Florida, Central America, tropical South America), Macroclinium (42; tropical America), Mesospinidium (8; western tropical South America)?, Miltonia (12; tropical America), Miltoniopsis (5; tropical America), Nohawilliamsia (1; N. pirarensis; Venezuela, Guyana, northern Brazil)?, Notylia (c 55; tropical America), Notyliopsis (2; Colombia), Oliveriana (6; northern South America), Oncidium (c 310; Florida and southwards to Chile and Argentina), Ornithocephalus (c 55; tropical America), Otoglossum (13; Central America, northern South America), Phymatidium (10; Brazil, Paraguay, Argentina), Platyrhiza (1; P. quadricolor; Brazil), Plectrophora (10; Trinidad, northeastern South America), Polyotidium (1; P. huebneri; Colombia, Venezuela, Brazil), Psychopsiella (1; P. limminghei; tropical America; in Psychopsis?), Psychopsis (4; tropical America; incl. Psychopsiella?), Pterostemma (3; Colombia, Ecuador), Quekettia (4; Trinidad, northern South America), Rauhiella (3; Brazil), Rhynchostele (17; tropical America), Rodriguezia (c 50; tropical America), Rossioglossum (9; Mexico, Central America, tropical South America), Sanderella (2; Brazil, Bolivia, Argentina), Saundersia (2; Brazil), Schunkea (1; S. vierlingii; Peru), Seegeriella (2; Bolivia), Solenidium (3; northern South America), Suarezia (1; S. ecuadorana; Ecuador), Sutrina (2; Peru, Bolivia), Systeloglossum (5; Costa Rica, Panama, Ecuador, Peru), Telipogon (c 205; Central America, tropical South America), Thysanoglossa (3; southeastern Brazil), Tolumnia (27; Florida, Mexico, Central America, the West Indies, northern South America), Trichocentrum (c 70; Florida, Mexico and southwards to Argentina), Trichoceros (10; northern South America), Trichopilia (c 45; tropical America), Trizeuxis (1; T. falcata; southern Central America, Trinidad, tropical South America), Vitekorchis (4; Central America, tropical South America), Warmingia (4; Costa Rica, Ecuador, Brazil, Paraguay, Argentina), Zelenkoa (1; Z. venusta; Panamá, Colombia to Peru), Zygostates (22; tropical South America). – Zygopetalinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 577, 588. 22 Jul 1926 [’Zygopetaleae’]. Aetheorhyncha (1; A. andreettae; Ecuador), Aganisia (4; tropical America), Batemannia (5; tropical South America), Benzingia (9; Costa Rica and southwards to Peru), Chaubardia (3; tropical South America), Chaubardiella (8; tropical America), Cheiradenia (1; C. cuspidata; northeastern South America), Chondrorhyncha (7; tropical America), Chondroscaphe (14; southern Central America, northwestern South America), Cochleanthes (4; tropical America), Cryptarrhena (3; tropical America), Daiotyla (4; Costa Rica to Colombia), Dichaea (c 120; tropical America), Echinorhyncha (5; Colombia, Ecuador), Euryblema (2; Panamá, Colombia), Galeottia (12; southern Mexico, Central America, tropical South America), Hoehneella (2; Brazil), Huntleya (14; tropical America), Ixyophora (5; tropical South America), Kefersteinia (c 70; tropical America), Koellensteinia (17; Central America, Puerto Rico, Trinidad, tropical South America), Neogardneria (1; N. murrayana; Brazil), Otostylis (4; Trinidad, tropical South America), Pabstia (5; Brazil), Paradisanthus (4; Brazil), Pescatoria (23; tropical America), Promenaea (18; Brazil), Stenia (22; Trinidad, northern South America), Stenotyla (9; southern Mexico, Central America), Vargasiella (2; Venezuela, Peru), Warczewiczella (11; Central America, Cuba, tropical South America), Warrea (3; tropical America), Warreella (2; Colombia), Warreopsis (4; southern Central America, northwestern South America), Zygopetalum (14; tropical America), Zygosepalum (8; tropical South America). – Eriopsidinae Szlach. in Fragm. Florist. Geobot., Suppl. 3: 97. 1995.Eriopsis (5; tropical America). – Maxillariinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [’Maxillarieae’].Anguloa (9; tropical South America), Bifrenaria (21; tropical America), Camaridium (c 140; Florida, Mexico, Central America, the West Indies, tropical South America southwards to Bolivia)?, Christensonella (10; Mexico, Central America, tropical South America)?, Chrysocycnis (4; Central America, Colombia, Ecuador)?, Cryptocentrum (22; Central America, tropical South America)?, Cyrtidiorchis (5; tropical South America)?, Guanchezia (1; G. maguirei; Venezuela), Heterotaxis (15; Florida, Mexico, Central America, the West Indies, tropical South America)?, Horvatia (1; H. andicola; Peru), Hylaeorchis (1; H. petiolaris; northwestern and northernmost South America)?, Inti (2; Central America, northern South America)?, Lycaste (32; tropical America), Mapinguari (4; Central America, northern South America)?, Maxillaria (c 660; Mexico, Central America, the West Indies, tropical South America), Maxillariella (c 45; Mexico, Central America, the West Indies, tropical South America)?, Mormolyca (c 25; Mexico, Central America, the West Indies, northern South America)?, Neomoorea (1; N. wallisii; Panamá to Ecuador), Nitidobulbon (3; Mexico, Central America, northwestern South America)?, Ornithidium (c 20; Mexico, Central America, the West Indies, tropical South America)?, Pityphyllum (7; northwestern South America)?, Rhetinantha (13; Mexico, Central America, tropical South America)?, Rudolfiella (6; tropical America), Sauvetrea (11; Central America, northern South America)?, Scuticaria (11; tropical South America), Sudamerlycaste (c 40; the West Indies, tropical South America), Teuscheria (7; northern South America), Trigonidium (c 25; tropical South America)?, Xylobium (c 30; tropical America). – Coeliopsidinae Szlach. in Fragm. Florist. Geobot., Suppl. 3: 97. 1995. Coeliopsis (1; C. hyacinthosma; Central America), Lycomormium (5; tropical South America), Peristeria (13; tropical America). – Stanhopeinae Benth. in J. Linn. Soc. London, Bot. 18: 288. 21 Feb 1881 [’Stanhopieae’]. Acineta (17; tropical America), Archivea (1; A. kewensis; Brazil)?, Braemia (1; B. vittata; northern South America), Cirrhaea (7; Brazil), Coryanthes (c 60; tropical America), Embreea (2; Colombia, Ecuador), Gongora (c 75; tropical America), Horichia (1; H. dressleri; Panamá), Houlletia (9; tropical America), Kegeliella (4; tropical America), Lacaena (2; Central America), Lueckelia (1; L. breviloba; Brazil), Lueddemannia (3; western tropical South America), Paphinia (12; tropical America), Polycycnis (c 15; tropical America), Schlimmia (7; northern Andes), Sievekingia (16; tropical America), Soterosanthus (1; S. shepheardii; tropical America), Stanhopea (c 60; tropical America), Trevoria (5; Colombia, Ecuador), Vasqueziella (1; V. boliviana; Bolivia). – Epidendreae Lindl., Collect. Bot.: 7, 16. Jan 1826. Agrostophyllinae Szlach. in Fragm. Florist. Geobot., Suppl. 3: 66. 1995.Agrostophyllum (c 100; the Seychelles and eastwards to western Pacific islands), Earina (7; western Pacific islands). – Calypsoinae Camus, Monogr. Orchid.: 376. 1908. Aplectrum (1; A. hyemale; Japan, North America), Calypso (1; C. bulbosa; temperate regions on the Northern Hemisphere), Changnienia (2; central and eastern China), Coelia (5; Central America, the West Indies), Corallorhiza (11; temperate regions on the Northern Hemisphere, with their highest diversity in North America), Cremastra (4; East Asia), Dactylostalix (1; D. ringens; Japan), Danxiaorchis (1; D. singchiana; Danxiashan in Guangdong), Ephippianthus (2; the Korean Peninsula, Japan, Sakhalin), Govenia (c 25; Florida, tropical America), Oreorchis (16; western Himalaya to Japan and Taiwan), Tipularia (7; Himalayas, Japan, eastern North America), Yoania (4; Himalayas, Japan, New Zealand). – Bletiinae Benth. in J. Linn. Soc. London, Bot. 18: 287. 21 Feb 1881 [’Bletieae’]. Basiphyllaea (7; Florida, the West Indies), Bletia (33; Florida, tropical America), Chysis (7–10; tropical America), Hexalectris (10; southern United States, Mexico).– Ponerinae Pfitzer, Entwurf. Anordn. Orch.: 101. Jan-Apr 1887 [’Ponereae’]. Helleriella (2; Central America), Isochilus (13; Mexico, Central America, Cuba, tropical South America), Nemaconia (6; tropical America), Ponera (2; tropical America). – Laeliinae Benth. in J. Linn. Soc. London, Bot. 18: 287. 21 Feb 1881 [’Laelieae’]. Acrorchis (1; A. roseola; Central America), Adamantinia (1; A. miltonioides; Serra do Sincorá in Brazil), Alamania (1; A. punicea; Mexico), Amoana (2; southern central Mexico)?, Arpophyllum (3; tropical America), Artorima (1; A. erubescens; Mexico), Barkeria (17; Central America), Brassavola (22; tropical America), Broughtonia (6; the Greater Antilles, the Bahamas), Cattleya (c 115; Central America, tropical South America), Caularthron (4; tropical America), Constantia (6; Brazil), Dimerandra (8; tropical America), Dinema (1; D. polybulbon; southern Mexico, Central America, Cuba, Jamaica), Domingoa (4; Central America, Cuba, Hispaniola, Venezuela), Encyclia (c 165; tropical America), Epidendrum (1.410–1.415; incl. Prosthechea?), Guarianthe (4; tropical America), Hagsatera (2; Mexico), Homalopetalum (8; tropical America), Isabelia (3; Brazil), Jacquiniella (12; tropical America), Laelia (23; tropical America; incl. Schomburgkia?), Schomburgkia (22–24; tropical America; in Laelia?), Leptotes (9; tropical South America), Loefgrenianthus (1; L. blanche-amesii; Brazil), Meiracyllium (2; southern Mexico, Central America), Microepidendrum (1; M. subulatifolium; Mexico), Myrmecophila (10; tropical America), Nidema (2; tropical America), Oestlundia (4; tropical America), Orleanesia (9; tropical South America), Prosthechea (c 120; tropical America; in Epidendrum?), Pseudolaelia (18; tropical America), Psychilis (14; the West Indies), Pygmaeorchis (2; Brazil), Quisqueya (4; the West Indies), Rhyncholaelia (2; Mexico, Central America), Scaphyglottis (c 70; Mexico, Central America, the West Indies, tropical South America), Tetramicra (13; the West Indies). – Pleurothallidinae Lindl. ex G. Don in R. Sweet, Hort. Brit., ed. 3: 636. late 1839 [’Pleurothalleae’]. Acianthera (c 120; Mexico to Chile), Anathallis (c 150; Mexico, Central America, the West Indies, tropical and subtropical South America), Andinia (13; tropical America), Barbosella (19; tropical America), Brachionidium (c 75; tropical America), Chamelophyton (1; C. kegelii; Venezuela, Guyana), Dilomilis (5; the West Indies to Brazil), Diodonopsis (5; mountain areas in tropical America), Draconanthes (2; the Andes, Ecuador), Dracula (125–130; tropical America), Dresslerella (13; Central America, northwestern South America), Dryadella (c 55; tropical America), Echinosepala (11; tropical America), Frondaria (1; F. caulescens; central Colombia to central Bolivia), Jostia (1; J. teaguei; Colombia, Ecuador, Peru)?, Kraenzlinella (9; southern Mexico, Central America, tropical South America), Lepanthes (c 1.085; tropical America), Lepanthopsis (c 45; tropical America), Masdevallia (c 590; mountain areas in tropical America), Myoxanthus (c 50; tropical America), Neocogniauxia (2; the West Indies), Octomeria (c 160; tropical America), Ophidion (4; Central America and southwards to the Andes; in Pleurothallis?)?, Pabstiella (c 30; southern Mexico, Central America, tropical South America), Phloeophila (11; Mexico, Central America, tropical South America), Platystele (c 100; tropical America), Pleurothallis (c 550; tropical America), Pleurothallopsis (18; Central America, tropical South America), Porroglossum (43; the Andes), Restrepia (53; tropical America), Restrepiella (2; Central America), Restrepiopsis (21; tropical America)?, Sansonia (2; tropical South America), Scaphosepalum (c 45; tropical America), Specklinia (c 135; Mexico, Central America, the West Indies, tropical South America), Stelis (c 880; tropical America), Teagueia (13; the Andes), Tomzanonia (1; T. filicina; Massif de la Hotte in Haiti), Trichosalpinx (c 110; tropical America), Trisetella (23; tropical America), Zootrophion (23; Central America, Jamaica, northern South America). – Collabieae Pfitzer, Entwurf. Anordn. Orch.: 100. Jan-Apr 1887 [’Collabiinae’]. Acanthephippium (13; tropical Asia and eastwards to Tonga), Ancistrochilus (2; tropical Africa), Ania (11; India, southern China, Southeast Asia, Malesia to New Guinea, the Solomon Islands, Queensland), Calanthe (c 215; tropical and subtropical regions in Madagascar, tropical Asia and eastwards to New Guinea, Australia and Tahiti, one species in tropical and southeastern Africa, one species in Central America and the West Indies), Cephalantheropsis (4; tropical Asia; in Calanthe?), Chrysoglossum (4; tropical Asia and eastwards to Samoa), Collabium (14; China to Malesia and Polynesia), Diglyphosa (3; tropical Asia), Eriodes (1; E. barbata; northeastern India to Vietnam), Gastrorchis (8; Madagascar; in Calanthe?), Hancockia (1; H. uniflora; East and Southeast Asia), Ipsea (3; India, Sri Lanka, Thailand), Nephelaphyllum (11; tropical Asia), Pachystoma (3; tropical Asia to China and western Pacific islands), Paraphaius (3; subtropical regions in East Asia), Phaius (c 45; tropical Africa, tropical Asia to southern China and eastern Queensland and northeastern New South Wales; in Calanthe?), Pilophyllum (1; P. villosum; Malesia), Plocoglottis (41; Malesia and eastwards to islands in the Pacific), Risleya (1; R. atropurpurea; Himalayas, western China), Spathoglottis (c 50; tropical Asia and eastwards to northeastern Queensland, Samoa and Niue), Tainia (23; tropical Asia to China and northeastern Queensland). – Podochileae Pfitzer, Entwurf. Anordn. Orch.: 101. Jan-Apr 1887 [’Podochilinae’]. Appendicula (c 145; tropical Asia and eastwards to Tonga), Ascidieria (8; southern Thailand, Malesia to the Philippines), Bryobium (8; Sri Lanka, southern China, Southeast Asia, Malesia to New Guinea, eastern Queensland, Melanesia), Callostylis (5; Assam, southern China, Southeast Asia, Java), Campanulorchis (5; Hainan, Southeast Asia, Malesia to New Guinea), Ceratostylis (c 150; tropical Asia and eastwards to western Pacific islands), Conchidium (10; India, southern China, Indochina, the Malay Peninsula), Cryptochilus (5; Himalayas), Dilochiopsis (1; D. scortechinii; the Malay Peninsula), Epiblastus (22; East Malesia and eastwards to Polynesia), Eria (c 240; tropical Asia), Mediocalcar (17; Sulawesi and eastwards to Samoa, with their largest diversity on New Guinea), Mycaranthes (36; Himalayas, southern China, Southeast Asia, New Guinea), Notheria (15; Sulawesi), Octarrhena (c 50; Sri Lanka, Malesia, New Caledonia, Polynesia), Oxystophyllum (36; southern China, Southeast Asia, Malesia to New Guinea, the Solomon Islands), Phreatia (c 210; northeastern India to Taiwan, New Guinea and tropical Australia), Pinalia (105; Himalayas, northeastern India to Burma, southern China, Thailand, Indochina, Malesia to New Guinea, Queensland, islands in the Pacific), Poaephyllum (6; Southeast Asia, Malesia), Podochilus (60–65; tropical Asia to southwestern China and western Pacific islands), Porpax (13; tropical Asia), Pseuderia (c 20; East Malesia, New Guinea to Fiji, Samoa and Micronesia), Rhynchophreatia (10; Malesia to New Guinea, the Solomon Islands, northern Australia, New Caledonia, Micronesia)?, Ridleyella (1; R. paniculata; New Guinea), Sarcostoma (5; the Malay Peninsula, Java, Sulawesi), Stolzia (15; tropical Africa), Thelasis (26; tropical Asia to southern China and the Solomon Islands), Trichotosia (c 80; India and eastwards to Vanuatu). – Vandeae Lindl., Orchid. Select.: 7, 14. Jan 1826. Polystachyinae Ridl., Mat. Flora Malay Penins. 1: 10. 1907 [’Polystachyae’]. Hederorkis (2; the Mascarene Islands), Imerinaea (1; I. madagascarica; Madagascar; in Polystachya?), Neobenthamia (1; N. gracilis; Uluguru and Nguru Mountains in Tanzania; in Polystachya?), Polystachya (c 235; tropical and southeastern Africa, Madagascar, tropical Asia eastwards to New Guinea and tropical Australia, tropical America). – Adrorhizinae Schltr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 573, 586. 22 Jul 1926 [’Adrorhizeae’]. Adrorhizon (1; A. purpurascens; Sri Lanka), Bromheadia (c 30; Sri Lanka and Burma to eastern Queensland), Sirhookera (2; southern India, Sri Lanka). – Aeridinae Pfitzer, Entwurf. Anordn. Orch.: 108. Jan-Apr 1887 [’Aerideae’]. Abdominea (1; A. minimiflora; Thailand, West and Central Malesia; in Tuberolabium?)?, Acampe (8; tropical and subtropical regions in the Old World), Adenoncos (17; Thailand, Vietnam, Malesia), Aerides (c 20; East Asia, tropical Asia to New Guinea), Amesiella (1; A. philippinensis; the Philippines), Arachnis (13; Southeast Asia and eastwards to the Solomon Islands), Armodorum (3; tropical Asia)?, Ascocentrum (12; Himalayas, Burma, Southeast Asia, West Malesia)?, Ascochilopsis (2; Thailand to Borneo)?, Ascochilus (6; Thailand to the Philippines)?, Ascoglossum (1; A. calopterum; New Guinea, the Solomon Islands)?, Biermannia (11; tropical Asia eastwards to Bali), Bogoria (4; Java, New Guinea), Brachypeza (10; Malesia), Calymmanthera (5; New Guinea), Ceratocentron (1; C. fesselii; the Philippines), Ceratochilus (1; C. biglandulosus; Java)?, Chamaeanthus (3; Thailand, Indochina, Malesia to Borneo), Chiloschista (c 20; India, southern China, Southeast Asia, Malesia to New Guinea, northern Australia, Fiji, Micronesia), Christensonia (1; C. vietnamica; Vietnam)?, Chroniochilus (4; Thailand to Borneo), Cleisocentron (6; Sikkim and eastwards to Vietnam and Borneo), Cleisomeria (2; Southeast Asia, Malesia), Cleisostoma (c 90; Nepal and eastwards to New Caledonia and Fiji), Cleisostomopsis (2; southern China, Indochina), Cordiglottis (7; southern Thailand, West Malesia)?, Cottonia (1; C. macrostachya; southern India, Sri Lanka), Cryptopylos (1; C. clausus; Southeast Asia, Sumatra), Deceptor (1; D. bidoupensis; Vietnam), Dimorphorchis (5; Borneo), Diplocentrum (2; India), Diploprora (2; tropical Asia), Dryadorchis (5; New Guinea), Drymoanthus (4; northeastern Queensland, New Zealand), Dyakia (1; D. hendersoniana; Borneo), Eclecticus (1; E. chungii; Thailand), Eparmatostigma (1; E. dives; Vietnam)?, Esmeralda (3; Himalayas, Assam, Tibet, southern China, northern Burma, northern Thailand, Hainan, Vietnam)?, Gastrochilus (c 55; Himalayas to Japan and Taiwan), Grosourdya (11; Southeast Asia, West Malesia), Gunnarella (9; New Caledonia), Holcoglossum (14; Southeast Asia to southern Japan and Taiwan), Hygrochilus (1; H. parishii; Assam, Bhutan, Yunnan, Burma, Southeast Asia)?, Hymenorchis (12; the Philippines to New Guinea, tropical Australia and New Caledonia), Jejewoodia (6; Borneo), Luisia (c 40; tropical Asia to Japan and Polynesia), Macropodanthus (8; Malesia), Malleola (c 30; southern China, Southeast Asia, Malesia to New Guinea, Melanesia)?, Megalotus (1; M. bifidus; the Philippines; in Saccolabium?)?, Micropera (21; tropical Asia to eastern Australia and New Caledonia), Microsaccus (12; tropical Asia), Microtatorchis (50–60; Southeast Asia, Taiwan, New Guinea, northern Australia, western Pacific islands)?, Mobilabium (1; M. hamatum; northeastern Queensland), Nothodoritis (1; N. zhejiangensis; Zhejiang in China)?, Omoea (2; Java, the Philippines), Ophioglossella (1; O. chrysostoma; New Guinea), Ornithochilus (3; Himalayas, China, Thailand)?, Papilionanthe (11; Himalayas and eastwards to Malesia), Papillilabium (1; P. beckleri; southeastern Queensland, northeastern New South Wales), Paraphalaenopsis (4; western Borneo), Pelatantheria (8; India to Taiwan and West Malesia), Pennilabium (15; Thailand to the Philippines), Peristeranthus (1; P. hillii; eastern Queensland, northeastern New South Wales), Phalaenopsis (c 70; Himalayas and eastwards to tropical Australia), Phragmorchis (1; P. teretifolia; the Philippines), Plectorrhiza (3; eastern Queensland, eastern New South Wales, estern Victoria, Lord Howe), Pomatocalpa (c 25; China to Malesia, tropical Australia and Polynesia), Porphyrodesme (3; New Guinea)?, Porrorhachis (2; Malesia), Pteroceras (27; tropical Asia), Renanthera (c 20; tropical Asia), Rhinerrhiza (1; R. divitiflora; New Guinea, eastern Queensland, northeastern New South Wales, the Solomon Islands), Rhinerrhizopsis (3; New Guinea, the Solomon Islands, northern Queensland), Rhynchogyna (3; Southeast Asia, Malesia), Rhynchostylis (3; tropical Asia), Robiquetia (c 45; tropical Asia and eastwards to Tonga), Saccolabiopsis (14; tropical Asia and eastwards to eastern Queensland and Fiji), Saccolabium (5; Sumatra, Java), Sedirea (2; China, the Korean Peninsula, Japan; in Aerides?)?, Santotomasia (1; S. wardiana; Luzon in the Philippines), Sarcanthopsis (5; tropical Asia and eastwards to northernmost Queensland and New Caledonia), Sarcochilus (25; estern Queensland to eastern Victoria, Tasmania, New Caledonia), Sarcoglyphis (12; tropical Asia), Sarcophyton (3; Southeast Asia), Schistotylus (1; eastern New South Wales), Schoenorchis (c 25; China, tropical Asia and eastwards to Samoa), Seidenfadenia (1; S. mitrata; Burma, Thailand), Seidenfadeniella (2; India, Sri Lanka), Singchia (1; S. malipoensis; southeastern Yunnan), Smithsonia (3; India), Smitinandia (3; tropical Asia), Spongiola (1; S. lohokii; Borneo), Stereochilus (7; India to Thailand and Vietnam), Taeniophyllum (c 185; tropical Africa and eastwards to Japan, Australia and Tahiti), Taprobanea (1; T. spathulata; southern India, Sri Lanka), Thrixspermum (c 160; Taiwan, tropical Asia and eastwards to West Pacific islands), Trachoma (14; Indochina, Malesia to New Guinea, Melanesia, northeastern Queensland and islands in the southwestern Pacific), Trichoglottis (c 70; Taiwan, tropical Asia and eastwards to Polynesia), Tuberolabium (11; tropical Asia), Uncifera (6; India and eastwards to West Malesia), Vanda (c 75; Himalayas to Australia), Vandopsis (4; Southeast Asia, Malesia), Ventricularia (2; Southeast Asia, West Malesia to Borneo), Xenikophyton (2; India)? – Angraecinae Summerh. in Kew Bull. 20: 188. 9 Sep 1966.Aerangis (c 60; tropical and southern Africa, Madagascar, the Mascarene Islands, Sri Lanka), Aeranthes (43; Zimbabwe, Madagascar, the Mascarene Islands), Ambrella (1; A. longituba; Madagascar), Ancistrorhynchus (17; tropical Africa), Angraecopsis (22; tropical Africa, Madagascar, the Mascarene Islands), Angraecum (c 200; tropical and southern Africa, Madagascar, the Mascarene Islands, Sri Lanka), Beclardia (2; the Mascarene Islands), Bolusiella (6; tropical and southern Africa), Calyptrochilum (2; tropical Africa, Brazil), Campylocentrum (c 65; Florida, tropical America), Cardiochilos (1; C. williamsonii; southern Tanzania, Malawi), Chauliodon (1; C. deflexicalcaratum; tropical West Africa), Cribbia (4; tropical Africa), Cryptopus (4; Madagascar, the Mascarene Islands), Cyrtorchis (18; tropical and southern Africa), Dendrophylax (14; Florida, the West Indies), Diaphananthe (33; tropical and subtropical Africa), Dinklageella (4; tropical West and Central Africa), Distylodon (1; D. comptum; Cameroon, Uganda), Eggelingia (3; tropical Africa), Erasanthe (1; E. henrici; Madagascar), Eurychone (2; tropical Africa), Jumellea (c 60; eastern tropical Africa, Madagascar, the Mascarene Islands, with their highest diversity in Madagascar), Lemurella (4; Madagascar), Lemurorchis (1; L. madagascariensis; Madagascar), Listrostachys (1; L. pertusa; tropical West and Central Africa), Margelliantha (6; northeastern Congo, Tanzania), Microcoelia (c 30; tropical and southern Africa, Madagascar), Mystacidium (10; southern tropical and southern Africa), Neobathiea (5; Madagascar), Nephrangis (2; tropical Africa), Oeonia (5; Madagascar, the Mascarene Islands), Oeoniella (2; Madagascar, the Comoro Islands, the Mascarene Islands, the Seychelles), Ossiculum (1; O. aurantiacum; Cameroun), Plectrelminthus (1; P. caudatus; tropical West Africa), Podangis (1; P. dactyloceras; tropical Africa), Rangaeris (6; tropical and southern Africa), Rhaesteria (1; R. eggelingii; Uganda), Rhipidoglossum (c 35; tropical and subtropical Africa), Sobennikoffia (4; the Mascarene Islands), Solenangis (8; tropical and southeastern Africa, Madagascar, the Mascarene Islands), Sphyrarhynchus (1; S. schliebenii; Tanzania), Summerhayesia (2; tropical Africa), Taeniorrhiza (1; T. gabonensis; Gabon), Triceratorhynchus (1; T. viridiflorus; tropical East Africa), Tridactyle (47; tropical and southern Africa), Ypsilopus (5; eastern and southern Africa). – Mainly tropical epiphytes; few species in temperate regions. Sometimes mycoheterotrophic holoparasites without leaves. CAM physiology often present. Roots sometimes with pneumathodes. Velamen usually present. Stegmata with silica bodies sometimes absent. Tilosomes often present. Leaves usually distichous, with conduplicate(-plicate) ptyxis, usually articulated, simetimes caducous or absent. Stomata often paracytic or tetracytic. Median outer stamen fertile and lateral inner stamens staminodial. Anther incumbent (by elongation of columella or by very early bending of anther; sometimes strongly convex), operculate and with ‘beak’. Pollinia usually clavate and hard (sometimes sectile); pollinium stipe developed from apical part of median stigmatic lobe. Pollen grains often inaperturate. Cotyledon usually very little developed. n = 5 and more. Plastid gene matK pseudogene. – Unplaced Epidendroideae Amparoa (2; southern Mexico, Central America), Devogelia (1; D. intonsa; the Moluccas, New Guinea). – Palmorchis may be sister to Neottieae or these two successive sister-groups to the remaining Epidendroideae. The position of Devogelia is unknown due to lack of fresh material.

Cladogram (simplified) of Orchidaceae based on DNA sequence data (Cameron 2004).

Cladogram of Apostasioideae, Cypripedioideae, and Vanilloideae based on DNA sequence data (Cameron 2004, 2009).

Cladogram of Orchidoideae, based on DNA sequence data (Cameron 2004).

Cladogram of a part of Epidendroideae, based on DNA sequence data (Cameron 2004).

Cladogram of a part of Epidendroideae, based on DNA sequence data (Cameron 2004).

RUSCACEAE M. Roem.

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Roemer, Handb. Allg. Bot. 3: 446. 1840 [’Ruscineae’], nom. cons.

Convallariaceae Horan., Prim. Lin. Syst. Nat.: 53. 2 Nov 1834 [’Convallariaceae s. Smilaceae’]; Ophiopogonaceae (Endl.) Meisn., Plant. Vasc. Gen., Tab. Diagn. 397, Comm. 308. 12-14 Feb 1842 [’Ophiopogoneae’]; Aspidistraceae (Endl.) Hassk., Cat. Hort. Bot. Bogor.: 32. Oct 1844 [’Aspidistreae’]; Eriospermaceae (Endl.) Lem. in V. V. D. d’Orbigny, Dict. Univ. Hist. Nat. 5: 402. 29 Mar 1845 [’Eriospermeae’]; Tupistraceae Schnizl., Iconogr. Fam. Regni Veg. 1: 72. Jan-Apr 1846 [’Tupistreae’]; Dracaenaceae Salisb., Gen. Plant.: 73. 15-31 Mai 1866 [’Dracaeneae’], nom. cons.; Peliosanthaceae Salisb., Gen. Plant.: 63. 15-31 Mai 1866 [’Peliosantheae’]; Platymetraceae Salisb., Gen. Plant.: 9. 15-31 Mai 1866 [‘Platymetreae’], nom. illeg.; Polygonataceae Salisb., Gen. Plant.: 63. 15-31 Mai 1866 [’Polygonateae’]; Sansevieriaceae Nakai in J. Jap. Bot. 12: 779. 1936; Nolinaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 226. 20 Jul 1943

Genera/species c 24/720–760

Distribution Temperate to tropical regions on the Northern Hemisphere southwards to West Malesia and Central America, with their largest diversity in East and Southeast Asia, North America, Mexico and South Africa.

Fossils Maianthemophyllum was described from Paleocene layers in Alberta, and Soleredera from the mid-Eocene of British Columbia.

Habit Usually bisexual (sometimes monoecious, polygamomonoecious, dioecious, or polygamodioecious), usually perennial herbs (sometimes suffrutices; rarely epiphytes; in Dracaena, Beaucarnea and Nolina frutescent or arborescent). Danae, Ruscus and Semele have green assimilating branches with ends modified into phyllocladia. Often xerophytic. Many species of Eriospermum and Comospermum have a hypocotyledonary tuber. Some species have lignotuber.

Vegetative anatomy Roots fibrous or fleshy (in some species of Ophiopogon swollen and carnose), often tuberous (tuber in Comospermum and Eriospermum developed from hypocotyl); velamen often present (sometimes multilayered). Danae, Ruscus and Semele have their phyllocladia and prophyll lateral or entirely fused with axillary shoot, together forming a compound phyllocladium. Phellogen? Secondary lateral growth usually absent (Dracaena, Beaucarnea, Nolana etc. with anomalous secondary lateral growth). Vessels present in roots and, in some species, in stem (at least rhizome) or leaves. Vessel elements in roots with scalariform or simple perforation plates (in stem only scalariform or absent); lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Vascular bundles sometimes amphivasal. Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Tanniniferous cells present in some genera. Dracaena secreting a blackish-red substance, hard and vitreous when oxidized and dry. Silica bodies absent. Mucilage cells with calciumoxalate raphides and styloids. Suberized cells with pseudo-raphides present in some genera.

Trichomes Hairs usually absent; leaf hairs in Eriospermum simple or compound; exotestal hairs in Eriospermum (and Comospermum?) unicellular, long, finally air-filled.

Leaves Usually alternate (spiral or distichous; rarely verticillate; in Polygonatum oppositifolium opposite), simple, entire, sometimes linear (in Danae, Ruscus and Semele reduced, scale-like, not photosynthesizing), often caducous (sometimes differentiated into pseudopetiole and pseudolamina), with conduplicate, supervolute or flat to curved ptyxis. Stipules absent (Peliosanthes sometimes with stipule-like outgrowths above leaf sheath); leaf sheath usually more or less well developed (sometimes absent). Leaf base in Eriospermum with filiform, bottle brush or feather duster like (sometimes bushy) appendages. Venation parallelodromous, sometimes acrodromous; midvein sometimes prominent; primary and secondary veins parallel. Stomata usually anomocytic (rarely paracytic or tetracytic), in Beaucarnea, Dasylirion and Nolina with oily guard cells. Cuticular wax crystalloids as parallel rodlets or platelets (Convallaria type) or non-orientated. Mesophyll often with mucilaginous idioblasts or cavities containing calciumoxalate raphides and/or single prismatic crystals. Leaf margin usually entire (in Dasylirion serrate-dentate).

Inflorescence Terminal or axillary, panicle, spike-, raceme- or head-like, or raceme or spike (flowers rarely solitary). In Semele and Ruscus one group (rarely two groups) of flowers on margin (umbel-like inflorescence) and on surface (raceme-like inflorescence) of a phyllocladium. Abaxial side of floral bracts sometimes with small nectaries.

Flowers Usually actinomorphic (sometimes slightly zygomorphic), usually small. Pedicel articulated, with pericladium. Hypogyny or half epigyny. Tepals (2–)3(–5)+(2–)3(–5), petaloid, caducous or marcescent, free or connate into a campanulate, urceolate or tubular perianth (in Aspidistra up to 13-lobate). Septal nectaries (infralocular?) usually present (sometimes absent; Convallaria without septal nectaries but with osmophores as tepaline oil glands; Tupistra with gynoecial nectaries). Extrafloral nectaries usually present at pedicel bases in Dracaena. Disc absent.

Androecium Stamens (2–)3(–6)+(2–)3(–6), usually as many (rarely twice as many) as tepals (outer staminal whorl sometimes staminodial or absent?). Filaments filiform to wide and flat, free or connate into a tube, free from or adnate to tepals or corona. Anthers basifixed or dorsifixed, in Dracaena and Eriospermum versatile, tetrasporangiate, usually introrse (sometimes latrorse or extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory, usually with binucleate cells. Female flowers often with staminodia (bisexual flowers and male flowers sometimes with three staminodia).

Pollen grains Microsporogenesis successive. Pollen grains monosulcate, monosulcoidate or inaperturate, shed as monads, usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate, or foveolate.

Gynoecium Pistil composed of (two or) three (to five) connate carpels. Ovary superior to semi-inferior, (unilocular to) trilocular (to quinquelocular). Style single, simple, filiform to stout, often with a stylar canal, or absent. Stigma capitate or trilobate (in ‘Aspidistra’ peltately widened), papillate or non-papillate, usually Wet type. Male flowers often with pistillodium.

Ovules Placentation usually axile (sometimes intrusively axile; sometimes parietal, basal, or apical). Ovules usually one to six (sometimes numerous) per carpel, anatropous, hemianatropous?, campylotropous or orthotropous, apotropous or epitropous, bitegmic, usually crassinucellar (sometimes tenuinucellar). Micropyle usually bistomal (in Danae, Ruscus, Semele, and Eriospermum endostomal). Outer integument two to eight cell layers thick. Inner integument two cell layers thick. Funicular obturator or ovary wall obturator often present (e.g. in Eriospermum). Parietal cell often formed from archesporial cell (absent in, e.g., Nolana, Beaucarnea and Dasylirion), and sometimes further dividing by periclinal divisions. Parietal tissue one to three (or four) cell layers thick. Nucellar cap usually absent (in, e.g., Dracaena, formed by periclinal cell divisions in megasporangial epidermis; no parietal cell formed). Megagametophyte usually monosporous, Polygonum type, or disporous, 8-nucleate, Allium type (in Maianthemum tetrasporous, 16-nucleate, Scilla or Drusa type). Chalazal end of megagametophyte in Ruscus contracted around antipodal cells and surrounded by a hypostase, a sclerotized lignified sheath. Synergids sometimes with a filiform apparatus. Antipodal cells usually not persistent (sometimes proliferating to five cells). Megagametophyte haustorium present in Dasylirion. Endosperm development usually helobial (sometimes nuclear). Endosperm haustoria? Embryogenesis in Eriospermum solanad, with megasporangial tissue enclosing chalazal parts of large embryo.

Fruit Usually a one- or several-seeded berry (in Tricalistra a drupe; in Comospermum, Eriospermum and Gonioscypha usually a loculicidal capsule; in Nolina, Beaucarnea, Calibanus and Dasylirion a samara; fruit in Liriope, Ophiopogon and Peliosanthes usually an irregularly dehiscing capsule).

Seeds Aril absent. Testa thin, degenerating (Liriope, Ophiopogon and Peliosanthes usually with sarcotesta; testa absent in Dracaena; testa in Eriospermum with long hairs). Exotesta without phytomelan layer, often with phlobaphene. Tegmen persistent, with thick cell walls, or collapsed. Perisperm usually not developed (in Eriospermum and possibly Comospermum thin, with oils and aleurone). Endosperm usually copious, with oils, aleurone and hemicellulose (starch absent); endosperm cell walls with pits or pores (endosperm absent from ripe seeds in Eriospermum). Embryo small to large (in Comospermum with coma), straight or curved, usually well differentiated, without chlorophyll. Cotyledon one, not photosynthesizing. Cotyledon hyperphyll compact, not assimilating. Hypocotyl internode short. Mesocotyl absent. Coleoptile present in Liriope, Ophiopogon and Peliosanthes. Radicula well developed, simple or branched, ephemeral or persistent. Germination? Seedling without ligule.

Cytology n = 5–7, 9, 10, 18–21 – Polyploidy occurring. Chromosomes 0.5–19 µm long. Karyotype bimodal (six large and 14 small chromosomes) in Danae, Ruscus, Semele and certain other species of Eriospermum. Adventitious polyembryony present (in, e.g., Maianthemum stellatum).

DNA The mitochondrial gene rpl2 is absent.

Phytochemistry Flavonols (e.g. 3-O-methyl-8-C-methylquercetin in, e.g., Nolina and Dasylirion, kaempferol in, e.g., Danae and Dracaena), cardenolide glycosides (Convallaria, Polygonatum, Speirantha), spirostanol glycosides (in Peliosanthes), steroidal saponins and sapogenins (e.g. diosgenin in Nolina, and aspidirin, a diosgenin-3-O-β-lycotetraoside, in Aspidistra), chelidonic acid, and azetidin-2-carboxylic acid (a non-protein amino acid) present. Fructans, polyacetate derived arthroquinones and chrysophanol present in roots of Ruscus. Ellagic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found.

Use Ornamental plants, medicinal plants, alcohol (sotol), textiles, basketry etc. (Nolina, Beaucarnea, Dasylirion, etc.).

Systematics Ruscaceae are sister-group to Asparagaceae.

Eriospermeae are sister to the remaining Ruscaceae.

Eriospermeae Endl. ex Meisn., Plant. Vasc. Gen.: Tab. Diagn. 397, 400, Comm. 304. 17-20 Aug 1842

1/c 100. Eriospermum (c 100; tropical and southern Africa, with their highest diversity in Western Cape). – Tuber of Eriospermum develop from hypocotyls. Leaves sometimes with various branched, peltate or stellate-haired etc. enations on adaxial surface. Pedicels non-articulated. Fruit a loculicidal capsule. Seeds often exposed early during development. Testa hairy. Embryo massive. Perisperm present. Endosperm absent. Embryo massive. Cotyledon unifacial, photosynthesizing. n = 20.

Rusceae Dumort., Anal. Fam. Plant.: 60. 1829 [‘Ruscineae’]

c 23/620–660. Disporopsis (6; Southeast Asia, the Philippines), Polygonatum (57; temperate regions on the Northern Hemisphere, with their highest diversity in southwestern China), Heteropolygonatum (3; China), Maianthemum (28; temperate regions on the Northern Hemisphere and southwards to Himalayas and Central America); Theropogon (1; T. pallidus; Himalayas), Comospermum (1; C. yedonense; Japan), Chrysodracon (6; the Hawaiian Islands), Dracaena (160–190; the Canary Islands, warm-temperate to tropical regions in the Old World, one species, D. americana, in Central America, one species, D. marginata, on Cuba), Danae (1; D. racemosa; southern Turkey, northwestern Syria, the Caspian area), Semele (1; S. androgyna; Macaronesia), Ruscus (6; western Europe, Macaronesia, the Mediterranean and eastwards to Iran); Peliosanthes (3–10; tropical Asia), Liriope (8; China, Japan, Vietnam), Ophiopogon (54; Himalayas to Japan, tropical Asia); Dasylirion (17; southwestern United States, Mexico), Nolina (23; southern United States, Mexico, Central America), Beaucarnea (5–10; Mexico, Belize, Guatemala); Speirantha (1; S. convallarioides; eastern China), Convallaria (1–3; C. majalis; temperate regions on the Northern Hemisphere), ’Aspidistra’ (c 100; East and Southeast Asia; paraphyletic), Reineckea (1; R. carnea; China, Japan), Rohdea (17; northern India, southern Himalayas, China, Japan, Indochina), Tupistra (c 20; northern India, southern Himalayas, southwestern and southern China, Southeast Asia, West Malesia, Ambon). – Temperate to tropical regions on the Northern Hemisphere southwards to West Malesia and Central America. – The clade [Peliosanthes+[Liriope+Ophiopogon]] has irregularly dehiscing capsule, sarcotesta and coleoptiles. Homoisoflavanones are present in Ophiopogon. Comospermum has two ovules per carpel, apotropous, tenuinucellate and the embryo has a coma (hair tuft).

Bayesian inference tree of Ruscaceae based on DNA sequence data (Kim & al. 2010).

TECOPHILAEACEAE Leyb.

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Leybold in Bonplandia 10: 370. Dec 1862 [‘Tecophilaeoceae’], nom. cons.

Androsynaceae Salisb., Gen. Plant.: 61. 15-31 Mai 1866 [‘Androsyneae’]; Cyanellaceae Salisb., Gen. Plant.: 46. 15-31 Mai 1866 [‘Cyanelleae’]; Conantheraceae (Endl. ex Meisn.) Endl. in L. K. G. Pfeiffer, Nomencl. Bot. 1: 839. 21 Feb 1873 [’Conanthereae’]; Cyanastraceae Engl. in Bot. Jahrb. Syst. 28: 357. 22 Mai 1900; Tecophilaeales Traub ex Reveal in Phytologia 74: 176. 25 Mar 1993; Walleriaceae (R. Dahlgren) H. Huber ex Takht. in Bot. Žurn. 79(12): 65. 7-28 Feb 1995, nom. cons.

Genera/species 8/25–26

Distribution Tropical and southern Africa, Madagascar, California, Chile.

Fossils Unknown.

Habit Bisexual, perennial herbs. Rhizome developed into a tuber or a corm (with basal innovation).

Vegetative anatomy Roots fibrous. Tuberous rhizome in most tunicate genera consisting of fibrous to membranous scale-like leaves, leaf sheaths, or vascular and fibre bundles (absent from Cyanastrum, Kabuyea and Walleria). Vascular bundles in one row, with or without sclerenchyma. 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 P2c, with cuneate protein crystals. Nodes? Calciumoxalate raphides usually present (absent in Cyanastrum and Kabuyea). Anthocyanin cells present in Cyanastrum and Kabuyea.

Trichomes Hairs usually absent (in Cyanella unicellular or short and multicellular, multiseriate; in Walleria scabrate).

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, often linear (in Cyanastrum and Kabuyea differentiated into pseudopetiole and pseudolamina), with ? ptyxis. Stipules absent; leaf sheath open? (sometimes absent). Venation parallelodromous (in Cyanastrum and Kabuyea pinnate-parallel or palmate-parallel), with distinct midvein. Stomata usually anomocytic (in, e.g., Cyanastrum indistinctly paracytic), adjacent cells with parallel divisions. Cuticular wax crystalloids absent (Cyanella). Mesophyll usually with cells containing calciumoxalate raphides. Mesophyll in Cyanastrum and Kabuyea with schizogenous secretory cavities (oil canals). Leaf margin entire.

Inflorescence Usually terminal, panicle or thyrsoid, often with raceme-like branches (flowers in Tecophilaea one or few, in Walleria solitary axillary, in Cyanastrum and Kabuyea in a simple raceme or pairwise). Some species of Cyanella have either monomorphic or dimorphic enantiostyly.

Flowers Actinomorphic or zygomorphic. Usually half epigyny (in Walleria almost hypogyny). Tepals 3+3, with imbricate aestivation, petaloid, free or connate at base into a tube; outer median tepal abaxial; outer tepals pointed. Coronae (enations), consisting of six small interstaminal appendages, present in Odontospermum and Zephyra. Septal nectaries usually situated over entire ovary (absent from or reduced to short septal slits in ovary apex in Cyanella, Tecophilaea and Walleria). Disc absent.

Androecium Stamens usually 3+3, all stamens fertile, or two (Zephyra) or three stamens (usually upper ones) staminodial; stamens in Cyanella usually dimorphic (heteranthery), with three small posterior and three large anterior stamens, or with five small posterior staminodial stamens and one large anterior fertile stamen (rarely all six stamens identical). Filaments subulate, usually free (rarely connate at base), adnate to tepals (epitepalous). Anthers usually free (in Conanthera connate), basifixed or subbasifixed (in Tecophilaea and Zephyra with basal caudate appendage), non-versatile, tetrasporangiate, poricidal (dehiscing by an apical or subapical pore), or introrse and dehiscing by short slits; connective sometimes apically prolonged. Tapetum secretory, with usually uni- or binucleate (in Cyanastrum multinucleate) cells. Staminodia sometimes two or three (rarely five). Enantiostyly occurring in some species of Cyanella.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate (sometimes trichotomosulcate), heteropolar, usually operculate (operculum absent in Cyanastrum and Kabuyea), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, usually foveolate, rugulate, granulate, in Kabuyea psilate or perforate.

Gynoecium Pistil composed of three, usually connate (in Cyanastrum free), antesepalous carpels; median carpel abaxial. Ovary usually semi-inferior (in Walleria almost superior), trilocular. Style single, simple, subulate, straight or curved (in Cyanastrum, Kabuyea and Walleria sunken and thus lateral on each carpel, gynobasic), with stylar canal. Stigma punctate or more or less trilobate, papillate, type? Pistillodium absent.

Ovules Placentation axile (in Cyanastrum and Kabuyea basal-axile?). Ovules usually four to c. 50 (in Cyanastrum, Kabuyea and Odontostomum two) per carpel, anatropous or campylotropous, bitegmic, crassinucellar. Micropyle endostomal or bistomal. Outer integument several cell layers thick. Inner integument two? cell layers thick. Funicular obturator usually present. Parietal cell formed from archesporial cell. Chalazal end of megasporangium usually well developed. Megagametophyte monosporous, Polygonum type. Endosperm development in Odontostomum and Cyanastrum probably helobial, in Cyanella nuclear. Endosperm haustorium in Cyanella chalazal. Embryogenesis?

Fruit A loculicidal capsule (in Walleria somewhat baccate; in Cyanastrum and Kabuyea? a schizocarp with three mericarps?).

Seeds Aril absent. Seed coat testal. Testa multilayered, thick-walled. Exotesta usually with phytomelan layer on epidermal cell walls (absent in Cyanastrum, Kabuyea? and Walleria), sometimes palisade, with thick cell walls. Endotesta? Tegmen? Perisperm not developed. Endosperm usually copious, thick-walled, with or without pits on surface, starchy? Endosperm absent in Cyanastrum, replaced by a swollen zone, chalazosperm, filled with starch grains, formed by cell divisions in chalazal tissue and consisting of loosely packed thin-walled cells (a similar starchy chalazal tissue present in Odontostomum). Embryo long or short, well differentiated, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll in Cyanella and Walleria with apex as an haustorium, a short tubular leaf sheath and a coleoptile. Hypocotyl absent in Cyanastrum. Radicula usually long (absent in Cyanastrum). Germination? Polyembryony present in Cyanella.

Cytology n = 8, 10–12, 14, 24 – Polyploidy and aneuploidy occurring. Chromosomes 2–4 µm long. Adventitious embryony present at least in Cyanella.

DNA

Phytochemistry Chelidonic acid present in Conanthera, and tuliposides in Tecophilaea. Steroidal saponins present in Conanthera? Flavonols, ellagic acid, proanthocyanidins, and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Tecophilaea (2; the Andes in Chile); Cyanastrum (3; tropical Africa), Kabuyea (1; K. hostifolia; Kenya, Tanzania); Odontostomum (1; O. hartwegii; California), Cyanella (7–8; Namibia, South Africa, with their highest diversity in Western Cape), Walleria (4; tropical and southern Africa, Madagascar), Zephyra (2; Chile), Conanthera (5; Chile).

Tecophilaeaceae are sister to Ixiolirion (Ixioliriaceae).

Tecophilaea in Chile is sister to the remaining Tecophilaeaceae and the African clade [Cyanastrum+Kabuyea] is successive sister-group to the rest. Odontostomum in California is sister to the African clade [Cyanella+Walleria] (Jesson & Barrett 2003).

Cladogram of Tecophilaeaceae based on DNA sequence data (Jesson & Barrett 2003).

THEMIDACEAE Salisb.

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Salisbury, Gen. Plant.: 84. 15-31 Mai 1866 [‘Themideae’]

Genera/species 12–15/c 65

Distribution Western North America to British Columbia, Mexico and Guatemala, with their largest diversity in the southwestern United States and Mexico.

Fossils Unknown.

Habit Bisexual, perennial herbs. Corm fibrous, rich in starch, and with basal innovation. Bulb absent.

Vegetative anatomy Young corm surrounded by dry leaf sheaths, cataphylls, of previous year, forming a membranous or fibrous tunica. Phellogen absent. Secondary lateral growth absent. Vessels present in roots. Vessel elements usually with scalariform or simple? perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Laticifers present at least in ‘Brodiaea’ and ‘Milla’. Mucilage cells? Calciumoxalate raphides present at least in Brodiaea.

Trichomes Hairs absent?

Leaves Alternate, simple, entire, linear, usually bifacial (in ‘Brodiaea’ unifacial), often fleshy, flattened, triangular or rounded in cross-section, with ? ptyxis. Stipules absent; leaf sheath closed. Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids? Mesophyll with or without cells containing calciumoxalate. Leaf margin entire.

Inflorescence Terminal, umbel-like, consisting of condensed helicoid monochasia (flowers rarely solitary), subtended by four or more, dry, membranous, early caducous bracts (spathae), not enclosing floral buds. Peduncle (scape) often long.

Flowers Actinomorphic. Pedicel often articulated. Hypogyny. Tepals 3+3, petaloid, usually connate in lower part into a tube (rarely free). Appendages, scales or corona often present between tepals and stamens (extrastaminal). Septal nectaries? Disc absent.

Androecium Stamens usually 3+3 (sometimes three fertile stamens, or three fertile stamens and three staminodia). Filaments usually subulate (sometimes flattened), often with basal, lateral, dorsal or apical appendages, free or connate (sometimes tubular), usually adnate to tepal (epitepalous); basal filament appendages in some species forming a nectary. Anthers basifixed or dorsifixed, usually versatile (not in ‘Brodiaea’ and ‘Dichelostemma’), tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia three or absent.

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

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular, often stipitate (with gynophore), in Bessera, ‘Milla’ and Petronymphe adnate to tepals by antesepalous flanges. Style single, simple, with stylar canal. Stigma capitate or trilobate (in ‘Brodiaea’ more or less squarrose), usually Dry (in Bloomeria Wet) type. Pistillodium absent.

Ovules Placentation axile. Ovules two or more per carpel, usually anatropous (rarely campylotropous), bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument three or four cell layers thick. Inner integument usually two or three (sometimes more than three) cell layers thick; inner integument and megasporangial base sometimes massive. Parietal tissue three or four cell layers thick. Megasporangial epidermal cells enlarged. Nucellar cap usually absent (sometimes present). Megagametophyte monosporous, Polygonum type (Brodiaea, Muilla). Endosperm development ab initio nuclear (‘Brodiaea’) or helobial (‘Muilla’, Triteleia). Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule.

Seeds Aril absent. Seed coat exotestal. Exotesta with thick phytomelan layer on epidermal cell walls. Mesotesta and endotesta compressed or collapsed. Tegmic cells usually strongly enlarged (not so in, e.g., Triteleia). Perisperm not developed. Endosperm copious, with oils and aleurone (starch absent). Embryo short, straight, without chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll? Hypocotyl internode? Coleoptile absent. Radicula persistent. Germination?

Cytology n = 5–12, 14–16, 18, 21, 24, 25, 27, 36, 42 – Polyploidy and aneuploidy frequently occurring.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Insufficiently known. Saponins present. Chlorogenic acid present at least in Brodiaea. Alkaloids and onion oils not found.

Use Ornamental plants.

SystematicsMilla’ (c 10; southwestern North America, Mexico, Guatemala; non-monophyletic), Bessera (3; California, Mexico), Petronymphe (1; P. decora; Mexico), Jaimehintonia (1; J. gypsophila; northeastern Mexico), Dandya (4; Mexico); Androstephium (3; western United States, Mexico), ‘Muilla’ (3–4; southwestern North America; non-monophyletic), Bloomeria (2; southwestern North America), Triteleia (18; western North America); Triteleiopsis (1; T. palmeri; Arizona, Baja California), ‘Brodiaea’ (c 15; western North America; non-monophyletic; in Dichelostemma?), ‘Dichelostemma’ (5; western North America; non-monophyletic; incl. Brodiaea?).

Themidaceae are sister to Hyacinthaceae.

Cladogram (simplified) of Themidaceae based on DNA sequence data (Pires & Sytsma 2002).

XANTHORRHOEACEAE Dumort.

( Back to Iridales )

Dumortier, Anal. Fam. Plant.: 60, 62, 103. 1829 [’Xanthorhaeaceae’, ’Xanthoraeaceae’, ’Xanthoarrheaceae’], nom. cons.

Xanthorrhoeales Takht. ex Reveal et Doweld in Novon 9: 552. 30 Dec 1999

Genera/species 1/c 30

Distribution Australia, Tasmania.

Fossils Unknown.

Habit Bisexual, evergreen, arborescent with aerial stem, or fruticose with subterranean stem. Stem thick, often tuberous, with leaves concentrated at apex.

Vegetative anatomy Roots contractile. Root cortex consisting of an ephemeral fleshy outer zone and a thickened inner zone; endodermal cells with U-shaped wall thickenings. Phellogen? Primary vascular stem tissue as scattered vascular bundles. Secondary lateral growth anomalous. Secondary thickening meristem dracaenoid. Vessels present at least in roots and leaves (sometimes in stem). Root vessel elements with simple perforation plates; lateral pits?; foliar and stem vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays? Axial parenchyma? Sieve tube plastids P2c type, with cuneate protein crystals. Nodes? Secretory ducts frequent, with yellow, brown or red acaroid resins containing essential oils. Silica bodies absent. Calciumoxalate raphides absent.

Trichomes Hairs?

Leaves Alternate (spiral), simple, entire, linear, unifacial, with ? ptyxis. Stipules absent; leaf sheath in young stems open (absent in older stems). Leaf bases thick and resiniferous, persistent as a dry coat around stem. Venation parallelodromous. Stomata paracytic, present in furrows. Cuticular wax crystalloids non-orientated and irregular. Mesophyll with cells containing calciumoxalate as single prismatic crystals. Secretory cavities and ducts with resin. Sclerenchymatic layers present inside epidermis. Leaf centre consisting of parenchyma with vascular bundles, some of them inverted. Epidermal cells with calciumoxalate as styloid-like crystals. Leaf margin entire.

Inflorescence Terminal, dense spike-like thyrse with alternate reduced cymes as partial inflorescences. Peduncle lignified. Each flower subtended by several bracts.

Flowers Actinomorphic. Pedicel not articulated. (Secondary) hypogyny. Tepals 3+3, sepaloid to bracteoid, persistent, free; outer tepals hard, dry; inner tepals membranous, almost petaloid. Septal nectaries well developed, infralocular. Disc absent.

Androecium Stamens 3+3. Filaments flattened, free from each other and from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate with widened sulcus, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary (secondarily) superior, trilocular. Style single, simple, subulate, narrowing at apex, with stylar canal. Stigma capitate, punctate or trilobate, probably Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules three to eight per carpel, anatropous, ascending or horizontal, bitegmic, crassinucellar. Micropyle endostomal. Outer integument approx. three cell layers thick. Inner integument two cell layers thick. Hypostase present. Megasporangium acute at apex. Megasporangial epidermal cells and parietal cells often dividing periclinally forming tissue up to six cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development helobial? Endosperm haustoria? Embryogenesis?

Fruit A lignified to cartilaginous loculicidal capsule.

Seeds Seeds flattened. Aril absent. Seed coat exotestal. Exotesta with phytomelan layer on epidermal cell walls. Endotesta? Tegmen collapsed, with inner cuticle of endotegmen present. Perisperm not developed. Endosperm copious, with oils and aleurone (starch absent), with relatively thick-walled cells containing small amounts of hemicellulose. Embryo straight to more or less curved, usually transversely oriented relative to seed axis, chlorophyll? Cotyledon one, not photosynthesizing. Cotyledon hyperphyll elongate. Hypocotyl internode short. Coleoptile absent. Germination phanerocotylar. First leaves distichous.

Cytology n = 11 – Karyotype bimodal.

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Flavonols (kaempferol, quercetin), flavans (e.g. 4’,5,7-trimethoxyflavan), chalcones, anthraquinones, polyacetate derived arthroquinones, and acaroid resins present (with, e.g., 2-hydroxy-4-methoxy acetophenone, 2-hydroxy-4,6-dimethoxy acetophenone, p-oxybenzoic aldehyde, xanthorrhein, citronellol, paeonol, cinnamoyl alcohol derivatives, p-coumaric acid and chrysophanic acid; in yellow resin also polymerized derivatives of cinnamic acids). Chelidonic acid? Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Ferulic acid absent from cell walls.

Use Wooden vessels, varnish (‘Grass Tree Gum’ and ‘Red Acaroid Gum’).

Systematics Xanthorrhoea (c 30; Australia, Tasmania).

Xanthorrhoea is sister-group to Hemerocallidaceae or, alternatively, to Asphodelaceae.

XERONEMATACEAE M. W. Chase, Rudall et M. F. Fay

( Back to Iridales )

Chase, Rudall et Fay in Kew Bull. 55: 869. 27 Dec 2000

Genera/species 1/2

Distribution New Caledonia, northernmost New Zealand.

Fossils Unknown.

Habit Bisexual, perennial herbs.

Vegetative anatomy 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 P2c type, with cuneate protein crystals. Nodes? Idioblasts with calciumoxalate raphides. Styloids absent.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, linear, equitant, unifacial, isobilaterally flattened, with ? ptyxis. Stipules absent; leaf sheath well developed. Venation parallelodromous. Stomata anomocytic? Cuticular wax crystalloids? Leaf margin entire.

Inflorescence Terminal, spike-like, unilateral.

Flowers Actinomorphic, directed upwards. Hypogyny. Tepals 3+3, petaloid, free. Septal nectaries well developed, branched. Disc absent?

Androecium Stamens 3+3. Filaments thick, subulate, free from each other and from tepals, long and strongly exserted. Anthers dorsifixed to subbasifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually monosulcate (rarely trichotomosulcate), boat-shaped, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, finely to coarsely reticulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, stipitate, trilocular. Style single, simple, solid. Stigma capitate, type? Pistillodium absent.

Ovules Placentation axile. Ovules several per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal? Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal cell? Parietal tissue? Nucellar cap formed? Megagametophyte monosporous, Polygonum type. Antipodal cells? Endosperm development helobial. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule.

Seeds Aril absent. Seeds echinate or papillate. Seed coat testal. Exotesta with phytomelan layer on epidermal cell walls. Endotesta? Tegmen collapsed. Perisperm not developed. Endosperm copious, without starch. Embryo straight?, chlorophyll? Cotyledon one. Cotyledon hyperphyll? Hypocotyl internode? Coleoptile absent. Germination?

Cytology n = 17, 18

DNA The mitochondrial gene rpl2 is absent (lost).

Phytochemistry Unknown.

Use Ornamental plants (X. callistemon).

Systematics Xeronema (2; X. moorei: New Caledonia; X. callistemon: Taranga and Poor Knights Island off northeastern New Zealand).

Xeronema is sister to the remaining Iridales (Hemerocallidaceae to Ruscaceae).


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