CARYOPHYLLIDAE Takht.

Takhtajan, Sist Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 144. 4 Feb 1967

[Berberidopsidales+Caryophyllales]

CARYOPHYLLALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 239. Jan-Apr 1820 [‘Caryophyllaceae’]

Caryophyllanae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 144. 4 Feb 1967

Fossils There are relatively few unambiguous fossils of Caryophyllales. Fossil flowers (e.g. Caryophylloflora paleogenica) with pantoporate pollen grains and campylotropous ovules, but with segmented ovary, have been found in Late Cretaceous (Turonian) and Eocene layers.

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes evergreen or deciduous trees, shrubs, suffrutices or lianas). Often leaf or stem succulents. Often halophytes. Often with spines. C₄ or CAM (crassulacean acid metabolism) physiologies often present as well as Kranz’ anatomy. Often mucilaginous. Sometimes carnivorous.

Vegetative anatomy Mycorrhiza absent in most groups (present in, i.a., Amaranthaceae and Nyctaginaceae). Root hair cells arranged in vertical rows. Phellogen epidermal, subepidermal, outer-cortical or pericyclic, or absent. Stem cortex often with two zones, outer with thick-walled fibrous cells, inner with thin-walled cells. Secondary lateral growth normal or anomalous (from concentric/successive cambia or from inner whorl of vascular bundles), or absent. Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes opposite or pseudoscalariform), usually simple (sometimes bordered) pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres (sometimes tracheids) with usually simple (sometimes bordered) pits, septate or non-septate (sometimes also vasicentric tracheids). Vestured pits sometimes present. Wood rays uniseriate or multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal, diffuse (sometimes diffuse-in-aggregates), or paratracheal, scanty vasicentric, aliform, winged-aliform, confluent, unilateral, or banded, or absent. Intraxylary phloem rarely present. Sieve tube plastids S₀, Ss, Pcs, P3c, P3cf, P3c’f, P3f, P3c’’f, P3cfs, or P3c’’fs type. Nodes usually 1:1, 1:3 or 3:3, unilacunar or with one or three leaf traces, or trilacunar with three traces (sometimes 5–9:5–9, multilacunar with five to nine traces). Parenchyma often with mucilaginous cells, often with sclereids. Wood with idioblasts containing sphaerites. Tanniniferous cells sometimes present. Silica bodies sometimes present in parenchyma cells. Calciumoxalate often present as druses, sphaerites, styloids, raphides, prismatic or acicular crystals, or crystal sand.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched (two-branched, T-shaped, dendritic, candelabra-shaped, stellate, fasciculate, lepidote, rosulate, barbed or vesicular), or absent; glandular hairs often frequent, multicellular, stalked or sessile (occasionally peltate-lepidote; sometimes secreting viscous mucilage).

Leaves Alternate (usually spiral, rarely distichous) or opposite (rarely verticillate), simple, usually entire (rarely lobed), often succulent, with conduplicate, supervolute, flat, involute or circinate ptyxis (sometimes scale-like or absent). Stipules usually absent (sometimes intrapetiolar); leaf sheath usually absent (leaf bases sometimes sheathingly connate). Petiole vascular bundle transection arcuate or annular, with peripheral ring of fibres. Venation pinnate or palmate, brochidodromous or parallelodromous (sometimes indistinct; leaves sometimes one-veined). Stomata usually paracytic, diacytic or anomocytic (sometimes anisocytic, cyclocytic, tetracytic or actinocytic, rarely brachyparacytic). Cuticular wax crystalloids as rodlets, threads or platelets, or absent. Leaf margin usually entire (rarely dentate, sinuate, serrate or glandular-serrate). Salt-secreting glands sometimes present.

Inflorescence Terminal or axillary, cymose combinations of dichasia and cincinni, thyrsoid, fasciculate, panicle, raceme-, spike- or head-like (flowers sometimes single). Floral prophylls (bracteoles) sometimes numerous, sometimes absent.

Flowers Usually actinomorphic (rarely zygomorphic). Hypanthium sometimes present. Usually hypogyny (rarely half epigyny). Sepals (one to) five (to 23), usually with imbricate (sometimes valvate, induplicate-valvate or open, rarely valvate-decussate, plicate or descending-cochlear) aestivation, free or more or less connate. Petals alt. petaloid staminodia (four or) five, with contorted or imbricate aestivation, free, or absent. Nectaries/nectariferous disc present at staminal bases or in tube formed by filament bases and petal bases or on inner side of receptacle, or nectary and disc absent.

Androecium Stamens (one to) five to more than 4.000 (staminal primordia usually five), in one or more whorls or in several groups (outer stamens often initiated in pairs), antetepalous when stamens isomerous relative to tepals; staminal development usually centrifugal. Filaments free or more or less connate, sometimes adnate to sepals or petals. Anthers basifixed or dorsifixed (sometimes latrorse), versatile or non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse (sometimes extrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores or pore-like slits); outer parietal cells developing directly into endothecium. Tapetum usually secretory (rarely amoeboid-periplasmodial). Female flowers often with staminodia (staminodia sometimes numerous in bisexual flowers). ‘Petaloids’ possibly in reality petaloid staminodia, developing simultaneously as or after androecium (not prior to); petaloid staminodia (’petals’) and ’antepetalous’ stamens possibly forming a developmental unit.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (rarely di- or tricolporate, or di- or tricolporoidate) or tetra-, hexa- or polypantoporate (sometimes tri- or hexarugate, rarely 4–12-colpate, spiraperturate, triporate, etc.), shed as monads, usually tricellular rarely bicellular) at dispersal. Exine tectate or semitectate (rarely intectate), with columellate infratectum, perforate, microperforate, punctate, punctitegillate or reticulate, scabrate, or spinulate.

Gynoecium Pistil composed of (one or) two to five (to numerous) free or connate carpels; closure of carpels sometimes delayed in at least Polygonaceae and in “the betalain clade” (Caryophyllales s.str.). Ovary superior, semi-inferior or inferior, unilocular to multilocular; ovary sometimes with subepidermal cell layer containing large amounts of calciumoxalate. Stylodia two to five, free or more or less connate, or style single, simple; style sometimes unifacial. Stigmas two to five, or stigma one, capitate or lobate, papillate, Dry type. Pistillodium usually absent (male flowers often with pistillodium).

Ovules Placentation axile, basal, basal-parietal, basal-lateral or free central (sometimes parietal-laminar, rarely apical or basal-laminar). Ovules one to numerous per carpel, campylotropous, hemianatropous or anacampylotropous (rarely anatropous, circinotropous or amphitropous), ascending, erect, horizontal or pendulous, apotropous or epitropous, usually bitegmic (rarely unitegmic), crassinucellar (rarely tenuinucellar). Placental or funicular obturator sometimes present. Micropyle usually endostomal (rarely bistomal). Funicle often with short hairs directed against micropyle. Megasporangium usually thin. Archespore unicellular to tricellular, only one developing further. Nucellar cap or nucellar beak often present. Apical cells of megasporangium often radially elongate, forming nucellar pad. Megagametophyte usually monosporous, Polygonum type (rarely disporous or tetrasporous, Allium, Adoxa,Endymion, Penaea, Drusa, Fritillaria, Chrysanthemum, Plumbago, or Plumbagella type). Synergids sometimes with a filiform apparatus. Antipodal cells three, ephemeral, sometimes with early degenerating nuclei. Chalazal caecum developed. Endosperm development ab initio usually nuclear (sometimes cellular). Endosperm haustoria chalazal or absent. Embryogenesis caryophyllad, chenopodiad or solanad (sometimes onagrad or asterad).

Fruit Usually a loculicidal and/or septicidal (rarely denticidal, circumscissile or valvate) capsule (sometimes a nut or an irregularly dehiscing capsule; rarely a berry, a berry-like fruit, a drupe, a schizocarp or a syncarp), sometimes with persistent calyx. Bracts and floral prophylls often partitioning in formation of dispersal unit.

Seeds Aril usually absent. Seed coat usually exotestal and/or endotegmic. Exotesta often tanniniferous; outer exotestal wall sometimes with stalactite-like processes. Endotesta sometimes thickened, often crystalliferous. Exotegmen? Endotegmen sometimes with rod-shaped thickenings in cell walls, often tanniniferous. Perisperm copious, starchy (with starch grains), surrounded by embryo, or not developed. Endosperm copious, sparse or absent. Embryo usually lateral-peripheral, curved around perisperm or straight (rarely cochleate or annular), with or without chlorophyll. Cotyledons usually two (rarely one or three). Germination phanerocotylar.

Cytology x = (4–)5–19

DNA Three large deletions present in plastid ORF2280. Intron absent from plastid gene rpl2. The plastid gene rpl23 is a pseudogene at least in the Caryophyllales analysed. I copy of nuclear gene RPB2 lost. Mitochondrial intron coxII.i3 sometimes lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavonol sulphates, flavone-C-glycosides (vitexin, isovitexin), flavones (e.g. luteolin), isoflavones, glycoflavones, cyanidin, delphinidin, catechines, anthocyanins or betalains (betacyanins, e.g. amaranthin, celosianin, betamin, phyllocactin, and betaxanthins), oleanolic acid derivatives, sterols, methylated and non-methylated ellagic acids, gallic acid, tannins, proanthocyanidins (prodelphinidins), mesembrine, tyramine alkaloids, phenethylamines, peyote alkaloids, indole alkaloids, tetrahydroisoquinoline alkaloids, acetogenic benzylisoquinoline alkaloids, naphthyl-isoquinoline alkaloids (e.g., ancistrocline, dioncophylline, michellamines), cyanogenic compounds (e.g. cyclopentenoid cyanogenic glycosides), betaine, triterpene saponins, anthraquinones, acetophenones, naphthoquinones and naphthoquinone derivatives (plumbagin, droserone, 5-O-methyl droserone, 7-methyljuglone, hydroxyserone; toxic naphthoquinones and their derivatives act as antimicrobial protectors and at least plumbagin as an insect ecdysis inhibitor e.g. in the pitchers of Nepenthes), simmondsin, fagopyrine, protofagopyrine, syringaresinol, oxalic acid, phytoferritin, phytoecdysones, cyclopeptides, and pinitol present. Ferulic acid often present in non-lignified cell walls. Inulin rarely present. Sulphated betalains present in the clade [Phytolaccaceae+ Petiveriaceae+Agdestis]. Benzylisoquinoline alkaloids and betalains both derived via shikimic acid (especially phenylalanine) pathway.

Systematics Caryophyllales are sister-group to Berberidopsidales.

Two strongly supported main clades can be discerned. One large monophyletic group comprises “the carnivorous clade” plus [[Tamaricaceae+Frankeniaceae]+[Polygonaceae+Plumbaginaceae]]. The second main clade includes [Rhabdodendraceae+[Simmondsiaceae+[Asteropeiaceae+Physenaceae]]] plus the core Caryophyllales in the strict sense.

Potential synapomorphies of the first main clade are according to Stevens (2001 onwards): presence of pit glands; endosperm starchy; and presence of acetogenic naphthoquinones. The first of the two subclades, “the carnivorous clade”, has the topology [[Droseraceae+Nepenthaceae]+[Drosophyllaceae+[Ancistrocladaceae+Dioncophyllaceae]]] and is characterized by the following potential synapomorphies: presence of vascularized multicellular glands; cymose inflorescence; corolla with contorted aestivation; extrorse anthers; unilocular ovary; and presence of plumbagin present. A clade identified in some analyses has the topology [Nepenthaceae+[Drosophyllaceae+[Ancistrocladaceae+Dioncophyllaceae]]] and the synapomorphies: presence of fibriform vessel elements; wood rays one or two cells wide; leaf vernation abaxially circinate; petiole vascular bundles surrounded by massive sclerenchymatous cylinder with embedded bundles; presence of wing bundles; and basifixed anthers. Ancistrocladaceae and Dioncophyllaceae share the following characters: climbing woody habit; phellogen deeply seated; absence of cortical vascular bundles; petiole with inverted vascular bundles in sclerenchyma cylinder; actinocyclocytic stomata; introrse anthers; and acetogenic naphthyl isoquinoline alkaloids biosynthesized from polyketides (not from aromatic amino acids).

The clade [[Frankeniaceae+Tamaricaceae]+[Plumbaginaceae+Polygonaceae]] is sometimes discerned and has the potential synapomorphies (Stevens 2001 onwards): vessel elements with minute lateral wall pits; outer and inner integuments two or three cell layers thick; exotestal seed coat; and presence of sulphated flavonols and ellagic acid. Frankeniaceae and Tamaricaceae share the characters: wood storied; halophytic habit with small leaves covered by salt-excreting glands; flowers small, tetra- to hexamerous; petals with basal adaxial appendages; exine not spinulate; median carpel abaxial; placentation usually parietal; loculicidal capsule; exotestal cells bulging or as hairs; presence of endosperm; presence of bisulphated flavonols; and absence of myricetin. Plumbaginaceae and Polygonaceae share the synapomorphies: wood storied; absence of successive cambia; presence of cortical and/or medullary vascular bundles; nodes 3:3; wide leaf bases; pollen grains usually starchy; median carpel adaxial; ovary unilocular; placentation basal; ovule one per carpel; fruit an anthocarp surrounded by accrescent calyx forming part of dispersal unit; seed coat indistinguished except persistent exotesta; loss of mitochondrial intron coxII.i3; and presence of O-methylflavonols, myricetin and quinones.

The second main clade has the topology [Rhabdodendraceae+[Simmondsiaceae+[[Asteropeiaceae+Physenaceae]+[Macarthuriaceae+[Microteaceae+[[Caryophyllaceae+[Achatocarpaceae+Amaranthaceae]]+[Stegnospermataceae to Cactaceae]]]]]]] and the potential synapomorphies (Stevens 2001 onwards): filament much shorter than anther; stylodia stigmatic (receptive) their entire length; ovules one or two per carpel; fruit single-seeded; and endosperm sparse. This clade minus Rhabdodendron is characterized by nodes 1:1 and absence of petals. Asteropeia and Physena lack successive cambia; have a vascular cylinder in the young stem; fibre tracheids; vasicentric tracheids; wood rays one or two cells wide; aliform-confluent axial parenchyma; latrorse anthers; and one-seeded fruit.

The core Caryophyllales – “the betalain clade” – comprise [Macarthuriaceae+[Microteaceae+[[Caryophyllaceae+[Achatocarpaceae+Amaranthaceae]]+[Stegnospermataceae to Cactaceae]]]]. Hence, they include the caryophyllids in the traditional strict sense. It is a group with very high bootstrap support and characterized by a large number of potential synapomorphies (Stevens 2001 onwards): herbaceous habit; absence of normal secondary lateral growth; CAM photosynthesis and C₄ photosynthesis frequently present; sieve tube plastids P-type, with a central angular protein crystal surrounded by a ring of protein filaments; absence of pericyclic fibres; cymose inflorescence; presence of adaxial nectaries on stamen bases; pollen grains tricellular at dispersal; exine with thin foot layer; median carpel adaxial; ovary unilocular; stigmas papillate; placentation free central or basal; ovules campylotropous; thickened exotestal and endotegmic cells; endotegmic cells with bar-like thickenings; perisperm well developed; absence of endosperm; starch grains clustered; embryo curved and periferal; cotyledons incumbent; absence of mitochondrial gene rps10; absence of plastid gene rpl2 intron; ferulic acid ester-linked to unlignified primary cell walls; presence of flavonols and O-methylated flavonols, quinones, usually betalains (chromoalkaloids) instead of anthocyanins, triterpenoid saponins and phytoferritin; and absence of tannins and myricetin. Usually phenylalanine-derived shikimic acid biosynthesis as starting point for synthesis of benzylisoquinoline alkaloids and betalains. An undifferentiated perianth evolved after the separation of the Rhabdodendron lineage, and a differentiated perianth may have originated at least nine times (Brockington & al. 2009): Asteropeia, Caryophyllaceae, Stegnosperma, species of Limeum, Corbichonia, Mesembryanthemoideae and Ruschioideae in Aizoaceae, Mirabilis in Nyctaginaceae, Glinus in Molluginaceae, Portulaca, Didiereaceae, Basellaceae, and Cactaceae.

The clade [Caryophyllaceae+[Achatocarpaceae+Amaranthaceae]] has the potential synapomorphies: stamens as many as tepals, antetepalous; a single ovule; parietal tissue approx. four cell layers thick; nucellar cap two to four cell layers thick; especially outer exotestal cell walls thick, with stalactite-like processes; mitochondrial genes rps1 and rps19 absent (lost); and often presence of phytoecdysteroids.

The lineages “above” Stegnosperma are characterized by apotropous ovules. “The globular inclusion clade” comprises Lophiocarpaceae to Cactaceae and posesses sieve tube plastids with globular crystalloids. The clade comprising Sarcobataceae, Nyctaginaceae, Agdestidaceae, Phytolaccaceae and Petiveriaceae often forms an unresolved polytomy (sometimes also including Gisekiaceae). It is characterized by a single usually basal ovule per carpel; similarities in ORF2280 sequence; and a 210 bp deletion in the plastid genome. They have usually (Sarcobatus?) a subepidermal phellogen; also paracytic stomata; and protein bodies in the nucleus. This clade minus Nyctaginaceae has racemose inflorescence and a baccate fruit.

The second large clade of “the globular inclusion lineage” comprises Molluginaceae to Cactaceae. The clade with the topology (Brockington & al. 2013) [[[Montiaceae+Halophytaceae]+[Didieraceae+Basellaceae]]+[Talinaceae+[Portulacaceae+[Anacampserotaceae+Cactaceae]]]] (Portulacineae, of Nyffeler & Eggli 2010; Cactineae of Ocamp & Columbus 2010) has the following potential synapomorphies, according to Stevens (2001 onwards): succulent leaves and/or stem; normal secondary lateral growth; phloem parenchyma cells with phytoferritin; stem epidermis with calciumoxalate crystals; presence of mucilage cells; leaves amphistomatic; median inner pair of floral prophylls enclosing flower; hypogyny; petaloid tepals; median tepal abaxial (opposite outer median floral prophyll); pollen grains pantocolpate; absence of funicular obturator; and a six bp deletion in plastid gene ndhF. The clade [Talinaceae+[Portulacaceae+[Anacampserotaceae+Cactaceae]]] is further characterized by columellae narrowed towards middle or expanded towards base, sometimes fused; pollen grains with granular internal surfaces; perforated foot layer; and very thin non-apertural endexine (Nowicke 1996). Halophytaceae, Basellaceae and Didiereaceae sometimes form a monophyletic group, with Halophytum sister to the remainder. Potential synapomorphies are ovary with a single basal ovule; and a single-seeded indehiscent fruit. Basellaceae may be sister-group to Didiereaceae or even nested within that clade (Didiereaceae and Basellaceae have often paracytic stomata). In this case, Halophytum may be sister to Didiereaceae including Basellaceae.

The monophyletic group [Talinaceae+[Portulacaceae+[Anacampserotaceae+Cactaceae]]] is supported by the potential synapomorphies (Stevens 2001 onwards): presence of mucilaginous cells; absence of pericyclic fibres; leaves with axillary uniseriate, biseriate or multiseriate hairs, bristles or scales; stomata parallelocytic (stoma with a lateral series of at least three alternating subsidiary cells increasing in size away from guard cells; also present in Montiaceae); pericarp two-layered; fruit covered by dry tepals; exocarp completely or almost caducous. Portulacaceae and Cactaceae may be sister-groups, according to Ocampo & Columbus (2010), using data mainly from non-coding plastid DNA. Portulaca and at least Pereskia share, e.g., a c. 500 bp deletion in rbcL, and non-lignified parenchyma cells are sometimes present in the wood in Portulaca and Cactaceae. On the other hand, Nyffeler & Eggli (2010), using sequence data from matK and ndhF, identified Anacampserotaceae and Cactaceae as sister-groups (they share among morphological features the character of numerous stamens). A special arrangement of testa cells along the dorsal juncture, presence of a dry aril and a central field type of cuticular ornamentation are characteristic features in several clades of Caryophyllales such as Cactaceae and Portulacaceae (Barthlott 1984).

A characteristic feature of the “ACPT clade” (Talinaceae, Portulaca, Anacampserotaceae and Cactaceae) are the non-vascularized hair-, bristle- or scale-like trichomes present at the nodes in the leaf axils (sometimes on internodes or on the lamina) (Ogburn & Edwards 2009). They arise from the epidermis and are persistent. The hair-like trichomes are either uniseriate or multiseriate (three or more cells in width). The bristle-like trichomes are wide, flat and multiseriate (up to more than 20 cells in width). The species in Anacampseros sect. Avonia possess wide scale-like trichomes which entirely surround the leaf distal to the subtending leaf; these scales are apically lignified. Bristle-like trichomes have been reported from species in Anacampserotaceae, whereas hair-like trichomes are present in Portulaca, Anacampserotaceae and Cactaceae. Membranous, often paired, scale-like trichomes are present in Talinum (Talinaceae). They have often been interpreted as homologous with the axillary trichomes, although they seem to be apices of vascularized prophylls subsequently often developing into leaves (Ogburn & Edwards 2009).

Problems concering homologies of the perianth parts are notorious in Caryophyllales. A uniseriate perianth is most probably plesiomorphic in Caryophyllales, and the uniseriate floral organs may be homologous (Brockington & al. 2009). A perianth may have originated by formation of homologous organs (perhaps in most lineages of Caryophyllales), or by differentiation of structures derived either from the androecium (in Corbichonia in Lophiocarpaceae, in the [Mesembryanthemoideae+Ruschioideae] clade of Aizoaceae, and in Molluginaceae) or from bracts (in Nyctaginaceae and in the “ACPT” clade), whereas ‘petaloid staminodia’ refer to floral parts that are anambiguously derived from the androecium. The terms ‘sepaloid tepals’ and ‘petaloid tepals’ are applied to floral organs (with quincuncial-imbricate aestivation) present in the core clade of Caryophyllales (Brockington & al. 2009). ‘Petaloid tepals’ in Caryophyllaceae, Stegnosperma, and Limeaceae have often been interpreted as modified stamens (e.g. Ronse De Craene 2007, 2008, etc.).

Phylogeny of Caryophyllales based on DNA sequence data (“core Caryophyllales” – Caryophyllineae: Brockington & al. 2013; “basal Caryophyllales” – Polygonineae: Brockington & al. 2009). ‘Hypertelis’ on this tree (according to Brockington & al. 2013) is now synonymous with Kewaceae, whereas the monospecific Hypertelis s. str. (H. spergulacea) is nested inside Molluginaceae> (Christenhusz & al. 2014).

Heimerl in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 16c: 174. Jan-Apr 1934, nom. cons.

Genera/species 2/c 11

Distribution California, Texas, Mexico to Paraguay and Argentina.

Fossils Unknown.

Habit Dioecious, evergreen trees or shrubs. Apices of short shoots often modified into spines.

Vegetative anatomy Phellogen ab initio superficial. Phelloderm with thick-walled sclereid cells. Secondary lateral growth normal (successive cambia absent). Vessel elements with simple perforation plates; lateral pits opposite or alternate, simple pits. Imperforate tracheary xylem elements libriform fibres with simple pits. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma (paratracheal) scanty vasicentric. Wood non-storied. Pericycle with sclerenchyma and stone cells. Phloem fibres present (scattered in older secondary phloem). Sieve tube plastids P3c’f type, with a single polygonal central protein crystal and a subperipheral dense ring of protein fibrils. Nodes? Tanniniferous cells often present. Calciumoxalate druses, sphaerites and prismatic crystals present (raphides and styloids absent).

Trichomes Hairs absent on older individuals; younger individuals with short hairs.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids as lobed platelets (often arranged in groups). Leaf margin entire.

Inflorescence Axillary, raceme-like, fasciculate or panicle. Foliar prophylls (bracteoles) absent in Achatocarpus.

Flowers Actinomorphic, small. Hypogyny. Tepals five (Achatocarpus) or usually four (rarely five; Phaulothamnus), with imbricate quincuncial (Achatocarpus) or decussate (Phaulothamnus) aestivation, sepaloid, free. Nectary absent. Disc absent.

Androecium Stamens ten to 20 (Achatocarpus) or twelve to 14 (Phaulothamnus). Filaments thin, free or connate at base. Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tetraporate to hexaporate or apertures more or less irregular and often little delimited, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, microperforate, scabrate (beset with microspinules) or coarsely granulate.

Gynoecium Pistil composed of two usually free carpels (sometimes slightly connate at base), collateral or superposed. Ovary superior, unilocular. Stylodia two, long, hairy and papillate, free or connate at base. Stigmas two, acute, hairy and papillate, type? Pistillodium absent.

Ovules Placentation basal. Ovule usually one (rarely two) per ovary, campylotropous, ascending, bitegmic, crassinucellar. Funicle present. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A usually one-seeded (occasionally two-seeded) berry.

Seeds Aril tiny (present at hilum). Testa? Outer exotestal wall with stalactite-shaped outgrowths? Tegmen? Perisperm copious, mealy. Endosperm rudimentary or absent. Embryo annular, peripheral, enclosing perisperm, chlorophyll? Cotyledons two. Germination?

Cytology n = ?

DNA

Phytochemistry Flavone-C-glycosides (vitexin, isovitexin) and tannins present. Anthocyanin and betalains? Ferulic acid in non-lignified cell-walls? Ellagic acid not found.

Use Medicinal plants.

Nakai in J. Jap. Bot. 18: 104. 10 Mar 1942

Genera/species 1/1

Distribution Southern United States to Nicaragua.

Fossils Unknown.

Habit Bisexual, evergreen, more or less lignified liana. Roots often napiform.

Vegetative anatomy Phellogen ab initio subepidermal. Secondary lateral growth anomalous (via concentric/successive cambia). Vascular bundles present as concentric cylinders in inner pericycle. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Imperforate tracheary xylem elements libriform fibres and tracheids with simple pits, septate or non-septate (also vasicentric tracheids). Wood rays absent. Axial parenchyma apotracheal, diffuse, or paratracheal. Wood non-storied. Tyloses sometimes present in vessels. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes? Parenchyma with idioblasts containing calciumoxalate as coarse raphide-like crystals.

Trichomes Hairs?

Leaves Alternate (spiral), simple, entire, with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole twisted at base. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids as rounded platelets. Idioblasts with calciumoxalate as coarse raphide-like crystals. Leaf margin entire.

Inflorescence Axillary, thyrsoid or panicle. Foliar prophylls (bracteoles) present.

Flowers Actinomorphic, small. Half epigyny. Tepals usually four (in terminal flowers sometimes five), with imbricate? aestivation, sepaloid, persistent, usually free (sometimes connate at base). Nectariferous disc narrow.

Androecium Stamens (twelve to) 15 to c. 30, in alternitepalous fascicles. Filaments filiform, usually connate at base (sometimes free), free from tepals, inserted at nectariferous disc. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate or perforate, scabrate, spinulate or smooth.

Gynoecium Pistil composed of (three or) four connate carpels. Ovary semi-inferior, ab initio (trilocular or) quadrilocular, later usually unilocular by degeneration of remaining locules. Style single, simple, short, cylindrical. Stigma (trilobate or) quadrilobate, recurved, papillate on ventral side, type? Pistillodium absent.

Ovules Placentation basal to axile. Ovule one per carpel, hemianatropous (or campylotropous?), bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue approx. two cell layers thick. Hypostase present. Nucellar cap massive. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A one-seeded unilocular achene with wings formed by persistent and accrescent dry tepals. Pericarp coriaceous, with reticulate pattern, adnate to seed coat.

Seeds Aril absent. Testa? Tegmen? Perisperm probably copious and nutritious. Endosperm poorly developed or absent. Embryo peripheral, curved around perisperm, well differentiated, chlorophyll? Cotyledons two. Germination?

Cytology n = 9 – Cell nuclei with protein bodies?

DNA 210 bp deletion present in plastid DNA?

Phytochemistry Betacyanins and betaxanthins present. Proanthocyanidins and alkaloids not found. Triterpenoid saponins? Free oxalates accumulated?

Use Medicinal plant, ornamental plant.

Martinov, Tekhno-Bot. Slovar: 15. 3 Aug 1820 [’Aizoonides’], nom. cons.

Mesembryanthemaceae Philib., Intr. Bot., ed. 2, 3: 268. 20 Dec 1801, [‘Mesembraceae’], nom. rejic.; Galeniaceae Raf. in Amer. J. Sci. 1: 376. Mai-Dec 1819; Mesembryaceae Dumort., Anal. Fam. Plant.: 37, 41. 1829 [‘Mesembryneae’]; Mesembryanthemales Link, Handbuch 2: 12. 4-11 Jul 1829 [‘Mesembrinae’]; Sesuviaceae Horan., Prim. Lin. Syst. Nat.: 83. 2 Nov 1834 [‘Sesuviaceae (Ficoideae)’]; Tetragoniaceae Lindl., Intr. Nat. Syst. Bot., ed. 2: 209. 13 Jun 1836, nom. cons.; Aizoales Boerl., Handl. Fl. Nederl. Ind. 1: li. 2 Aug 1890; Aizoineae Doweld, Tent. Syst. Plant. Vasc.: xli. 23 Dec 2001

Genera/species 124/1.675–1.695

Distribution Arid tropical and subtropical regions including western and southern Australia, with their highest diversity in southern and southwestern Africa. Mesembryanthemoideae and, above all, Ruschioideae dominate much of the succulent vegetation in the Karroo areas of South Africa, where they constitute more than half the number of species and more than 90% of the biomass.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious or dioecious), perennial or annual herbs (rarely climbing), suffrutices or shrubs. Usually leaf succulents (sometimes stem succulents, rarely root succulents; some species are almost entirely subterranean). Almost all representatives are xerophytes; some species are halophytes. C₄ or CAM (rarely C₃) physiology present. Kranz’ anatomy present in some species. Usually mucilaginous.

Vegetative anatomy Mycorrhiza usually absent. Kranz’ anatomy present in some species. Phellogen ab initio inner-cortical or endodermal, or absent. Primary medullary strands wide. Medullary wide-band tracheid cells frequent, also present in foliar tissue other than mid-vein; bands narrow yet very tall, cell lumen in places very narrow. Endodermis significant. Secondary lateral growth normal or sometimes anomalous (via concentric/successive cambia) or absent. Vessel elements usually with simple perforation plates; lateral pits alternate? Imperforate tracheary xylem elements libriform fibres with bordered pits? (in at least one species of Ruschia also vasicentric tracheids). Wood rays usually absent (present in Tetragonia). Axial parenchyma paratracheal? (in Tetragonia also vasicentric). Wood elements often partially storied. Sieve element plastids P3cf type, with a central globular protein crystal surrounded by protein filaments. Nodes 1:1, unilacunar with one leaf trace, or 3:3, trilacunar with three traces. Tanniniferous idioblasts often present in water-storing tissue. Epidermis in many species with densely spaced water-storing vesicular idioblasts with wide base. Calciumoxalate crystals often present.

Trichomes Hairs usually unicellular or multicellular and uniseriate (rarely stipitate, two-branched or stellate), often vesicular, or absent.

Leaves Usually opposite (often pairwise fused; sometimes alternate), simple, entire, often cuboidal, conical, cylindrical or prismatic, with usually curved or flat ptyxis. Stipules usually absent (petiole base in Sesuvioideae with stipule-like appendages); leaf bases often membranous and sheathingly connate around stem (sometimes entirely connate). Petiole vascular bundles forming cortical reticulum. Venation pinnate or palmate, usually indistinct, with sunken main veins. Stomata anomocytic, paracytic or anisocytic. Cuticular wax crystalloids as rodlets (often terete), threads or platelets. Mesophyll sometimes with idioblasts containing calciumoxalate raphides. Epidermis of upper side of lamina often with numerous water-storing wide-based vesicular idioblasts. Outer epidermal cell walls often with calciumoxalate crystals. Leaf margin entire, dentate or serrate.

Inflorescence Terminal (often seemingly axillary), cymose (sometimes capitate), or flowers usually solitary terminal. Floral prophylls (bracteoles) usually large, often foliaceous.

Flowers Actinomorphic, often large. Hypanthium usually present. Hypogyny, half epigyny or epigyny. Tepals (three to) five (to eight), in one whorl, with usually imbricate quincuncial (rarely valvate) aestivation, usually sepaloid (adaxial tepals sometimes petaloid), persistent, connate at base, often with subapical abaxial appendage. Nectary usually continuous, annular on adaxial side of perianth-stamen tube (nectaries in Mesembryanthemoideae partially tubular). Disc present or absent.

Androecium Stamens usually numerous (to more than 2.000; sometimes four, five, eight, or ten), in one or more whorls or in three to nine groups; staminal primordia five, alternitepalous, or as annular meristem. Outer whorls usually consisting of petaloid staminodia, inner whorls consisting of fertile stamens, median whorls often intermediary. Filaments free or connate (all or in three to nine groups), free from tepals. Anthers dorsifixed, often versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with usually trinucleate (rarely septanucleate) cells. Staminodia petaloid, extrastaminal, usually numerous (absent in some genera).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (rarely tetracolpate or tricolporoidate), shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, punctate or anulopunctate, spinulate (rarely reticulate or rugate).

Gynoecium Pistil composed of (one or) two to five (to numerous) connate carpels. Ovary superior to inferior, (unilocular or) bilocular to quinquelocular (to multilocular), usually entirely septate (septa in Acrosanthes incomplete). Stylodia usually free (rarely connate). Stigmatic areas adaxial, papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation usually axile (in early stage, primary; rarely basal or apical) to parietal (in later stage, secondary). Ovules one to numerous per carpel (in Acrosanthes a single basal ovule per carpel), campylotropous (hemianatropous?) to anacampylotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument two or three cell layers thick. Inner integument two cell layers thick. Obturator placental. Parietal tissue approx. three cell layer thick. Nucellar cap micropylar. Apical cells of megasporangium often radially elongate. Megagametophyte usually monosporous, Polygonum type (rarely Endymion, Penaea, Drusa, or Adoxa type). Endosperm development ab initio nuclear. Endosperm haustoria chalazal? Embryogenesis caryophyllad or solanad.

Fruit Usually a loculicidal capsule (sometimes a nut, rarely a pyxidium, a berry or a schizocarp; in Gunniopsis a septicidal capsule), usually hydrochastic, dehiscing in moist weather, when septa and other tissues swell (septal keels, reaching from central axis to valve apices).

Seeds Aril sometimes present (surrounding seed in Sesuvioideae). Exotesta palisade or tangentially elongate. Endotesta and tegmen usually crushed. Perisperm copious, starchy. Endosperm sparse or absent. Suspensor massive, usually biseriate to multiseriate. Embryo peripheral, curved around perisperm, without chlorophyll. Cotyledons two. Radicula dorsal. Germination phanerocotylar.

Cytology x = 8 (Sesuvioideae, Aizooideae); x = 9 (Mesembryanthemoideae, Ruschioideae) – Polyploidy frequently occurring.

DNA Intron absent from plastid gene rpoC1 (at least in Delospera and Faucaria). Plastid gene infA transferred to nucleus (Mesembryanthemum; pseudogene present in plastid genome).

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, condensed tannins, betacyanins, betaxanthins, mesembrine alkaloids (Phyllobolus etc.), and ferulic acid (in non-lignified cell walls) present. C-glycosylflavonoids, ellagic acid and saponins not found. Some species oxidize free oxalates.

Use Ornamental plants, vegetables (Tetragonia tetragonioides), fruits (Carpobrotus edulis), stabilization of soil (Carpobrotus etc.).

Cladogram (simplified) of Aizoaceae based on morphology and DNA sequence data (Chesselet & al. 2002, etc.).

de Jussieu, Gen. Plant.: 87. 4 Aug 1789 [’Amaranthi’], nom. cons.

Atriplicaceae Juss., Gen. Plant.: 83. 4 Aug 1789 [’Atriplices’]; Chenopodiaceae Vent., Tabl. Règne Vég. 2: 253. 5 Mai 1799 [’Chenopodae’], nom. cons.; Amaranthales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 240. Jan-Apr 1820 [‘Amaranthaceae’]; Celosiaceae Martinov, Tekhno-Bot. Slovar: 117. 3 Aug 1820 [’Celosiae’]; Chenopodiales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 240. Jan-Apr 1820 [‘Chenopodeae’]; Salicorniaceae Martinov, Tekhno-Bot. Slovar: 558. 3 Aug 1820 [’Salicorniae’]; Amaranthopsida Horan., Prim. Lin. Syst. Nat.: 58. 2 Nov 1834 [’Amaranthoideae’]; Betaceae Burnett, Outl. Bot.: 591, 1091, 1142. Feb 1835; Achyranthaceae Raf., Fl. Tellur. 3: 35. Nov-Dec 1837 [’Achyranthidia’]; Gomphrenaceae Raf., Fl. Tellur. 3: 38. Nov-Dec 1837 [’Gomphrenidia’]; Polycnemaceae Menge, Cat. Plat. Grudent. Gedan.: 161. 1839 [’Polycneminae’]; Salsolaceae Menge, Cat. Plant. Grudent. Gedan.: 165. 1839 [’Salsolinae’]; Spinaciaceae Menge, Cat. Plant. Grudent. Gedan.: 166. 1839 [’Spinacinae’]; Chenopodiineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1272, 1274. 1846; Atriplicales Horan., Char. Ess. Fam.: 63. 30 Jun 1847 [‘Atriplicastra s. Curvembryae’]; Deeringiaceae J. Agardh, Theoria Syst. Plant.: 369. Apr-Sep 1858 [’Deeringieae’]; Blitaceae Post et Kuntze, Lex. Gen. Phan.: 637, 710. 20-30 Nov 1903; Dysphaniaceae (Pax) Pax in Bot. Jahrb. Syst. 61: 230. 15 Jun 1927, nom. cons.

Genera/species 176/2.000–2.075

Distribution Cosmopolitan except polar areas, with their highest diversity in saline, arid and semiarid areas.

Fossils Fossil pollen (e.g. Chenopodipollis) are known from Late Cretaceous (Maastrichtian), Paleocene and Late Eocene layers in North America.

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, dioecious, androdioecious, rarely polygamomonoecious), perennial, biennial or annual herbs, evergreen or deciduous suffrutices or shrubs (rarely trees or lianas), sometimes with spines. Often leaf or stem succulents. C₄ plants with c. 17 different types of foliar anatomy. Many species are halophytes or xerophytes.

Vegetative anatomy Mycorrhiza usually absent (sometimes with vesicular-arbuscular mycorrhiza). Kranz’ anatomy present in numerous species. Phellogen usually superficial or pericyclic. Stem collenchyma well developed. Cortical and/or medullary bundles usually present. Primary medullary strands usually wide. Endodermis usually significant. Secondary lateral growth usually anomalous (polycyclic, anomalous secondary vascular bundles from concentric cambia; not in Polycnemoideae) or absent. Pericyclic fibres few or absent. Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes opposite), simple or bordered pits. Imperforate tracheary xylem elements libriform fibres (with cell nuclei) with simple or (reduced) bordered pits, non-septate (also vasicentric tracheids). Wood rays usually absent (sometimes uniseriate or multiseriate, homocellular or heterocellular). Axial parenchyma paratracheal scanty, aliform, winged-aliform, confluent, vasicentric, or banded. Vessel elements, fibres and/or parenchyma sometimes partially or entirely storied. Intraxylary (concentric or diffuse) phloem present. Sieve tube plastids P3cf type, without a central protein crystal, with circular peripheral protein fibrils (sometimes with starch grains). Nodes 1:1 or 1:3, unilacunar with one or three leaf traces (sometimes 1:5, unilacunar with five traces), often swollen. Heartwood sometimes with gum-like substances. Calciumoxalate usually as crystal sand, druses or prismatic crystals (raphides and styloids absent).

Trichomes Hair types unicellular to multicellular, uniseriate, T-shaped or many-armed (also malpighiaceous hairs), dendritic, stellate, candelabra-shaped, fasciculate, lepidote, capitate and/or vesicular, often with salt-storing apical cell; glandular hairs often present.

Leaves Alternate (spiral) or opposite, simple, usually entire (sometimes lobed; in some stem succulents reduced), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate; leaves sometimes one-veined. Stomata usually anomocytic (sometimes paracytic, diacytic or anisocytic). Cuticular wax crystalloids as platelets (Chenopodioideae, Salsoloideae etc.) or without platelets (Amaranthoideae etc.; cuticular wax crystalloids absent in Dysphania). Domatia present in many species. Hydathodes sometimes present. Leaf margin serrate, sinuate, crenate or entire.

Inflorescence Terminal or axillary, cymose, head-, raceme- or spike-like, thyrsoid, fasciculate or panicle (flowers sometimes solitary, axillary). Lateral flowers of dichasial partial inflorescences sometimes sterile and modified into scales, spines, bristles or hairs. Bracts often membranous or coloured. Floral prophylls (bracteoles) often petaloid (sometimes membranous). Extrafloral nectaries rarely present (Iresine).

Flowers Actinomorphic, small. Usually hypogyny (rarely half epigyny). Tepals (one or) three to five (to eight), with usually imbricate (rarely valvate) aestivation, sepaloid, often fleshy, often with tuberculate, spinulose or wing-like outgrowths, persistent, free or connate at base, or rudimentary or absent. Nectary absent. Disc (sometimes lobate) present in some species.

Androecium Stamens (one or) three to five (to nine), usually as many as tepals, antetepalous, or absent. Filaments free or connate in lower part (sometimes entirely) into a tube, often adnate to tepals (epitepalous); tuberculate, scale-like or fringed staminodium-like lobes, pseudostaminodia, often present between stamens and usually fused with filaments. Anthers usually dorsifixed, often versatile, usually tetrasporangiate (sometimes disporangiate), usually introrse (rarely extrorse), longicidal (dehiscing by longitudinal slits); connective sometimes with apical appendage (sometimes vesicular and coloured); anther wall development monocotyledonous. Tapetum usually secretory, with binucleate or multinucleate cells (sometimes amoeboid-periplasmodial). Staminodia one to five, often petaloid, or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains polypantoporate, shed as monads, usually tricellular (rarely bicellular) at dispersal. Pollen grains often with starch. Exine tectate or semitectate, with columellate infratectum, microperforate, punctate or reticulate (in Gomphrenoideae metareticulate), spinulate or tubuliferous.

Gynoecium Pistil composed of (one or) two or three (to six) connate carpels (median carpel sometimes abaxial); when two carpels then usually transverse. Ovary usually superior (rarely semi-inferior), unilocular. Style single, simple, or stylodia two or three, long or short, more or less connate. Stigma one, capitate (simple or penicillate), or stigmas two or three (to six), narrowly elongate, papillate, Dry type, often persistent. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation usually basal (sometimes free central or apical). Ovule usually one (ovules sometimes few or many) per ovary, campylotropous, anacampylotropous or circinotropous (rarely amphitropous), erect to pendulous, bitegmic, crassinucellar. Micropyle usually endostomal (rarely bistomal). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Nucellar beak present. Apical cells of megasporangium often radially elongate. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type). Synergids sometimes with a filiform apparatus. Antipodal cells usually persistent. Chalazal caecum developed. Endosperm development ab initio nuclear. Endosperm haustorium chalazal or absent. Embryogenesis chenopodiad or solanad.

Fruit Usually a nut (often a utriculus or an achene) or an irregularly dehiscent capsule (often a pyxidium, rarely a berry, a baccaceous fruit or a drupe; adjacent ovaries sometimes fusing and forming a syncarp), often surrounded by more or less fleshy perianth forming an anthocarp; persistent and often accrescent bracts and floral prophylls sometimes forming parts of dispersal unit.

Seeds Aril usually absent. Seed coat usually exotestal-endotegmic. Outer exotestal cell walls with stalactite-shaped appendages. Outer exotegmic cell walls often tanniniferous. Endotegmen often thickened and lignified, tanniniferous. Perisperm usually copious and starchy (rarely absent), usually surrounded by embryo. Endosperm sparse or absent. Embryo curved or annular around perisperm (rarely straight; in Salsoloideae spirally twisted), well differentiated, with or without chlorophyll. Cotyledons usually two (rarely three). Radicula dorsal. Germination phanerocotylar.

Cytology n = 6–36 – Polyploidy frequently occurring.

DNA Intron absent from plastid gene rpl2. Deletion of 300 bp in plastid IR in many clades. Plastid genome in at least Chenopodium and Atriplex with 6 kb inversion.

Phytochemistry Flavonols (kaempferol, quercetin), 6-7-methylene-dioxyflavonols, isoflavones, flavonol sulphates, betacyanins (e.g. amaranthin, celosianin and betamin or phyllocactin), betaxanthins, isoquinoline alkaloids and other alkaloids (particularly in Salsoleae), triterpene saponins, cyanogenic compounds, betaine, anthraquinones, and sterols present. Ferulic acid present in non-lignified cell walls. Ellagic acid and proanthocyanidins not found. Nitrate or free oxalates accumulated in many species.

Use Ornamental plants, vegetables (Beta vulgaris, Spinacia oleracea, Chenopodium quinoa, Atriplex hortensis, etc.), sugar (Beta vulgaris var. altissima), forage-plants (Beta vulgaris, Atriplex, Chenopodium, Rhagodia, Cornulaca), glass production (Salicornia, Salsola, etc.), timber, wood carving, medicinal plants (Dysphania etc.).

Systematics Amaranthaceae are sister-group to Achatocarpaceae or to the clade [Caryophyllaceae+Achatocarpaceae].

A plausible topology is, according to Kadereit & al. (2006) and Kadereit & Freitag (2011): [Polycnemoideae+[[Amaranthoideae+[Betoideae+[Corispermoideae+Chenopodioideae]]]+[[Suaedoideae+Salicornioideae]+[‘Hammada’tamariscifolia+[Salsoloideae+Camphorosmoideae]]]]].

Polycnemoideae Raf., Fl. Tellur. 3: 44. Nov-Dec 1837 [‘Polycnemides’]

4/14. Hemichroa (1; H. pentandra; Australia, Tasmania), Surreya (2; S. diandra, S. mesembryanthema; Australia), Nitrophila (5; N. atacamensis, N. australis, N. mexicana, N. mohavensis, N. occidentalis; western United States, northwestern Mexico, Chile, Argentina), Polycnemum (6; P. arvense, P. fontanesii, P. heuffelii, P. majus, P. perenne, P. verrucosum; central, southern and eastern Europe, the Mediterranean, northewesternmost Africa, Central Asia). – Europe, the Mediterranean, Central Asia, Australia, western United States, northwestern Mexico. Normal secondary lateral growth present. Flowers unisexual. Filaments connate at base. Anthers in Polycnemum monothecal (disporangiate). Pollen grains smooth or spinulose. – Polycnemoideae were sister to all other Amaranthaceae in one matK/trnK analysis (Müller & Borsch 2005), but nested deeply within Amaranthaceae in analyses by Kadereit & Freitag (2011). Polycnemum is sister to the clade [Nitrophila+[Hemichroa+Surreya]] (Masson & Kadereit 2013).

[[Amaranthoideae+[Betoideae+[Corispermoideae+Chenopodioideae]]]+[[Suaedoideae+Salicornioideae]+[‘Hammada’tamariscifolia+[Salsoloideae+Camphorosmoideae]]]]

[Amaranthoideae+[Betoideae+[Corispermoideae+Chenopodioideae]]]

Amaranthoideae Burnett, Outlines Bot.: 591, 593, 1091, 1142. Feb 1835 [‘Amarantidae’] (under construction)

76/800–830. Bosea (3; B. yervamora: the Canary Islands; B. cypria: Cyprus; B. amherstiana: western Himalayas); Charpentiera (6; C. densiflora, C. elliptica, C. obovata, C. ovata, C. tomentosa: the Hawaiian Islands; C. australis: Tubuai in the Austral Islands). – Amarantheae Rchb., Fl. Germ. Excurs. 2(2): 575, 583. 1832. Amaranthus (c 50; cosmopolitan), Chamissoa (2; C. acuminata, C. altissima; warmer regions in North to South America). – Celosieae Fenzl in S. F. L. Endlicher, Gen. Plant.: 303. Oct 1837. Celosia (c 50; warmer regions in North to South America), Deeringia (12; tropical and subtropical regions in the Old World), Henonia (1; H. scoparia; Madagascar), Hermbstaedtia (c 15; tropical and southern Africa except the Cape provinces), Pleuropetalum (3; P. pleiogynum, P. sprucei: Mexico to Peru; P. darwinii: the Galápagos Islands). – Pleuropetalum has racemose inflorescence, five tepals, eight stamens developed in pairs, filaments connate at base, pistil composed of five or six connate carpels, basal placenta with several ovules, fruit ab initio fleshy, and n = 8 or 9. – Aerveae Fenzl in S. F. L. Endlicher, Gen. Plant.: 302. Oct 1837. Aerva (c 20; tropical and subtropical regions in the Old World), Nothosaerva (1; N. brachiata; tropical Africa from Senegal to Ethiopia and northern Somalia, south to southern Africa, Mauritius, Pakistan, India, Sri Lanka, Burma, Mauritius), Ptilotus (100–110; drier regions in Australia, one species, P. conicus, also on Flores and Timor). – Gomphreneae Fenzl in S. F. L. Endlicher, Gen. Plant.: 301. Oct 1837. Irenella (1; I. chrysotricha; Ecuador), Iresine (35–40; tropical West Africa, southern Japan, tropical and subtropical America), Woehleria (1; W. serpyllifolia; Cuba); Alternanthera (130–140; tropical and subtropical regions on both hemispheres, with their largest diversity in tropical America), Pedersenia (9–10; tropical America), Tidestromia (6; T. carnosa, T. lanuginosa, T. rhizomatosa, T. suffruticosa, T. tenella, T. valdesiana; southwestern United States, Mexico); Blutaparon (4; B. portulacoides, B. rigidum [extinct], B. vermiculare, B. wrightii; tropical West Africa, Ryukyu Islands, tropical and subtropical America), Froelichia (15; tropical and subtropical America, the Galápagos Islands), Froelichiella (1; F. grisea; Brazil), ‘Gomphrena’ (130–135; tropical and subtropical regions in North to South America; polyphyletic), Gossypianthus (2; G. lanuginosus, G. tenuiflorus; United States, Mexico, Central America), Guilleminea (8; Central America), Hebanthodes (1; H. peruviana; Peru), Lecosia (2; L. formicarum, L. oppositifolia; southeastern Brazil), Lithophila (2; L. radicata, L. subscaposa; the Galápagos Islands), Pfaffia (c 30; tropical South America), Pseudogomphrena (1; P. scandens; Brazil), Pseudoplantago (2; P. bisteriliflora, P. friesii; Venezuela, Argentina), Quaternella (3; Q. confusa, Q. ephedroides, Q. glabratoides, Brazil), Xerosiphon (2; X. angustiflorus, X. aphyllus; tropical South America). – Pseudoplantago has cuboid or prismatic pollen grains. – Achyrantheae Fenzl in S. F. L. Endlicher, Gen. Plant.: 302. Oct 1837. Achyranthes (12; tropical and subtropical regions in the Old World; incl. Nototrichum), Nototrichium (3; N. divaricatum, N. humile, N. sandwicense; the Hawaiian Islands; in Achyranthes), Calicorema (2; C. capitata, C. squarrosa; tropical Africa and southwards to Namibia and Northern Cape), Cyathula (c 30; tropical regions in Africa, Asia, America and the Pacific), Pandiaka (13; tropical and southern Africa), Psilotrichum (16; tropical regions in the Old World), Pupalia (4; P. grandiflora, P. lappacea, P. micrantha, P. robbechii; tropical regions in the Old World), Sericostachys (1; S. scandens; tropical Africa). – Unplaced Amaranthoideae Achyropsis (6; A. avicularis, A. filifolia, A. fruticulosa, A. gracilis, A. laniceps, A. leptostachya; tropical and southern Africa), Allmania (1; A. nodiflora; tropical Asia), Allmaniopsis (1; A. fruticulosa; eastern Kenya), Arthraerua (1; A. leubnitziae; coast of Namibia), Centema (2; C. angolensis, C. subfusca; southern tropical Africa to South Africa), Centemopsis (13; tropical Africa), Centrostachys (1; C. aquatica; northern Africa, India, Java), Chionothrix (2; C. latifolia, C. somalensis; Somalia), Dasysphaera (4; D. alternifolia, D. hyposericea, D. robecchii, D. tomentosa; East Africa), Digera (1; D. muricata; tropical regions in the Old World), Eriostylos (1; E. stefaninii; Somalia), Herbstia (1; H. brasiliana; Brazil), Indobanalia (1; I. thyrsiflora; southwestern India), Kyphocarpa (3; K. angustifolia, K. cruciata, K. trichinoides; southern tropical and southern Africa), Lagrezia (9; Madagascar, Indian Ocean islands), Lecosia (2; L. formicarum, L. oppositifolia; southeastern Brazil), Leucosphaera (1; L. bainesii; Angola, Namibia, Botswana, northwestern Northern Cape), Lopriorea (1; L. ruspolii; East Africa), Marcelliopsis (3; M. denudata, M. splendens, M. welwitschii; Namibia), Mechowia (1; M. grandiflora; southern tropical Africa), Nelsia (3; N. angolensis, N. quadrangula, N. tropidogyna; Angola, southern Africa), Neocentema (2; N. alternifolia: Tanzania; N. robecchii: Somalia), Nyssanthes (2; N. diffusa, N. erecta; eastern Queensland, eastern New South Wales), Omegandra (1; O. kanisii; northern Australia), Pleuropterantha (3; P. revoilii, P. thulinii, P. undulatifolia; northeastern tropical Africa), Polyrhabda (1; P. atriplicifolia; Somalia), Pseudosericocoma (1; P. pungens; southwestern and southern Africa), Psilotrichopsis (1; P. curtisii; Thailand, the Malay Peninsula), Rosifax (1; R. sabuletorum; Somalia), Saltia (1; S. papposa; southern Arabian Peninsula), Sericocoma (3; S. avolans, S. heterochiton, S. pungens; tropical Africa to Namibia, Northern, Western and Eastern Cape), Sericocomopsis (2; S. hildebrandtii, S. pallida; tropical East Africa), Sericorema (2; S. remotiflora, S. sericea; southern Africa), Stilbanthus (1; S. scandens; the Himalayas), Trichuriella (1; T. monsoniae; southern and southeastern Asia), Volkensinia (1; V. prostrata; East Africa). – Subcosmopolitan with their largest diversity in tropical and subtropical America and Africa. – Bosea and Charpentiera are successive sisters to the remaining Amaranthoideae. Gomphreneae, with bilocular anthers and metareticulate exine, are sister-group to Achyrantheae.

[Betoideae+[Corispermoideae+Chenopodioideae]]

Betoideae Ulbr. in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 16c: 455. 28 Apr 1934

6/17. Acroglochin (2; A. obtusifolia, A. persicarioides; Central and East Asia, the Himalayas); Beta (9; Europe, the Mediterranean, temperate Asia), Aphanisma (1; A. blitoides; California, northwestern Mexico), Hablitzia (1; H. tamnoides; the Caucasus), Oreobliton (1; O. thesioides; Algeria, Tunisia), Patellifolia (3; P. patellaris, P. procumbens, P. webbiana; Macaronesia). – Europe, Macaronesia, the Mediterranean region, North Africa, southwestern Asia, southwestern North America (Aphanisma). Annual to perennial herbs (in Hablitzia twining) or suffrutices. Tepals usually five (in Aphanisma three). Fruit a pyxidium. – Betoideae are sister-group to the clade [Corispermoideae+Chenopodioideae]. Acroglochin is sister to the remaining Betoideae.

[Corispermoideae+Chenopodioideae]

Corispermoideae Raf., Fl. Tellur. 3: 45. Nov-Dec 1837 [‘Corispermides’]

3/c 80. Agriophyllum (6; A. lateriflorum, A. latifolium, A. minus, A. montasirii, A. paletzkianum, A. squarrosum; Europe, western Asia to Central Asia), Anthochlamys (5; A. afghanica, A. multinervis, A. polygaloides, A. tjanschanica, A. turcomanica; southwestern and Central Asia), Corispermum (c 70; temperate regions of Europe and Asia). – Eurasia. Annual herbs. C₃ metabolism and corispermoid non-Kranz’ leaf anatomy present. Hairs on young shoots usually dendritic (not in Anthochlamys). Floral prophylls (bracteoles) absent. Tepals one to five, membranous, caducous (sometimes absent). Perisperm copious. Embryo erect. – Corispermoideae are sister-group to Chenopodioideae.

Chenopodioideae Burnett, Outlines Bot.: 591, 1091, 1142. Feb 1835 [‘Chenopodidae’]

23/460–470. Dysphanieae Pax in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. III, 1b: 69, 92. Mai 1889. Cycloloma (1; C. atriplicifolium; western and central North America), Dysphania (>40; warmer regions on both hemispheres), Suckleya (1; S. suckleyana; Rocky Mountains), Teloxys (1; T. vagans; Mongolia). – Axyrideae G. Kadereit et Sukhor. in Amer. J. Bot. 97: 1682. Oct 2010. Axyris (6; A. amaranthoides, A. caucasica, A. hybrida, A. koreana, A. prostrata, A. sphaerosperma; Central Asia to the Korean Peninsula), Ceratocarpus (1; C. arenarius; eastern Europe, southwestern to Central Asia), Krascheninnikovia (3; K. ceratoides, K. ewersmanniana, K. fruticulosa; the Mediterranean, temperate Asia, western North America). – Anserineae Dumort., Fl. Belg.: 20. 1827. Blitum (12; Europe, northern and eastern Asia, Australia, Canada, United States, temperate South America), Spinacia (3; S. oleracea, S. tetrandra, S. turkestanica; southwestern Asia, North Africa). – Atripliceae Duby, Bot. Gall. 1: 394. 12-14 Apr 1828. Lipandra (1; L. polysperma; Europe, the Mediterranean), Oxybasis (5; O. chenopodioides, O. glauca, O. macrosperma, O. rubra, O. urbica; temperate regions in Europe and Asia, the Mediterranean), Chenopodiastrum (5; C. badachschanicum, C. coronopus, C. hybridum, C. murale, C. simplex; temperate regions on the Northern Hemisphere), Atriplex (250–260; cosmopolitan), Microgynoecium (1; M. tibeticum; Tibet), Archiatriplex (1; A. nanpinensis; Sichuan), Exomis (1; E. microphylla; Namibia, Northern, Western and Eastern Cape, Free State), Extriplex (2; E. californica, E. joaquinana; California), Grayia (3; G. arizonica, G. plummeri, G. spinosa; western United States), Holmbergia (1; H. tweedii; Uruguay, Paraguay, Argentina), Manochlamys (1; M. albicans; Namibia, Northern and Western Cape), Proatriplex (1; P. pleiantha; the Navajo Basin in southwestern United States), Stutzia (2; S. covillei, S. dioica; western United States), Chenopodium (c 115; cosmopolitan). – Chenopodioideae are sister-group to Corispermoideae. The Atriplex clade is sister-group to Chenopodium with high support.

[[Suaedoideae+Salicornioideae]+[‘Hammada’tamariscifolia+[Salsoloideae+Camphorosmoideae]]]

[Suaedoideae+Salicornioideae]

Suaedioideae Ulbr. in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 16c: 445, 554. 28 Apr 1934

2/75–80. Bienertia (3; B. cycloptera, B. kossinskyi, B. sinuspersici; southeastern European Russia and Kazakhstan to western Central Asia), Suaeda (70–75; cosmopolitan). – Subcosmopolitan. Annual or perennial herbs or shrubs. Leaf-succulents, halophytes. C₃ or C₄ metabolism with or without Kranz’ anatomy. Flowers in axillary cymes. Tepals five, connate at base. Perisperm usually absent. Embryo spiral. – Suaedioideae are sister-group to Salicornioideae.

Salicornioideae Luerss., Hand. Syst. Bot. 2: 547. Nov 1880

12/105–110. Heterostachys (2; H. olivascens: Argentina; H. ritteriana: Central America, Hispaniola, South America), Allenrolfea (3; A. occidentalis: southwestern United States, Mexico; A. patagonica, A. vaginata: Argentina), Halopeplis (3; H. amplexicaulis: the Mediterranean, North Africa; H. perfoliata: northeastern Africa, the Arabian Peninsula to Pakistan; H. pygmaea: the Caucasus and Iraq to Central Asia and Xinjiang), Halostachys (1; H. belangeriana; southeastern Russia to Central Asia), Halocnemum (2; H. strobilaceum, H. yurdakulolii; central Mediterranean to Central Asia), Kalidium (6; K. capsicum, K. cuspidatum, K. foliatum, K. gracile, K. schrenkianum, K. sinicum; the Mediterranean to Central Asia), Microcnemum (1; M. coralloides; the Mediterranean to the Caucasus), Arthrocaulon (2; A. franzii, A. macrostachyum; the Mediterranean, Macaronesia, northwestern and northeastern Africa, southwestern Asia), Arthroceras (1; A. subterminale; California, northern Mexico), Tecticornia (44; Australia, one species, T. indica, along coasts of eastern Africa, southern Asia and northern Australia), Mangleticornia (1; M. ecuadorensis; coasts of southern Ecuador and northern Peru), Salicornia (40–45; Europe, the Mediterranean, Africa, Asia, Australia, America). – Subcosmopolitan. Annual or perennial herbs or shrubs. Stem-succulents, halophytes. Usually with C₃ metabolism (one species of Tecticornia with C₄ metabolism). Flowers in spike-like thyrse. Embryo spiral. – Salicornioideae are sister to Suaedioideae.

[‘Hammada’ tamariscifolia+[Salsoloideae+Camphorosmoideae]]

Salsoloideae Raf., Fl. Tellur. 3: 45. Nov-Dec 1837 [‘Salsoloides’]

c 35–37/c 370. Salsoleae Dumort., Fl. Belg.: 22. 1827. Sympegma (1; S. regelii; Central Asia), Kali (13; Europe, the Mediterranean, temperate Asia), Traganum (2; T. moquinii, T. nudatum; the Canary Islands, eastern Mediterranean, North Africa), Xylosalsola (4; X. arbuscula, X. chiwensis, X. paletzkiana, X. richteri; Central Asia), Turania (4; T. androssowii, T. aperta, T. deserticola, T. sogdiana; Iran, Afghanistan, Central Asia, northwestern China), Halothamnus (c 20; the Middle East to Afghanistan, Somalia, Central Asia, northwestern China), Noaea (5; N. cadmea, N. major, N. minuta, N. mucronata, N. regelii; arid and semi-arid regions in eastern Europe to Central Asia), Oreosalsola (9; Iran, Afghanistan, Central Asia), ‘Salsola’ arbusculiformis (Iran, Central Asia, Xinjiang), ‘Salsola’ (c 50; Europe, Macaronesia, the Mediterranean, North Africa, southwestern and Central Asia; polyphyletic), Salsola divaricata (the Canary Islands), Haloxylon (2; H. ammodendron, H. persicum; the Mediterranean, North Africa, southwestern and Central Asia, Mongolia, western and northwestern China), Cornulaca (7; C. alaschanica, C. amblyacantha, C. aucheri, C. ehrenbergii, C. korshinskyi, C. monacantha, C. setifera; Egypt to Central Asia), Horaninovia (2; H. minor, H. ulicina; southwestern and Central Asia), Hammada (3; H. griffithii, H. leptoclada, H. wakhanica; southwestern and Central Asia), Halogeton (4; H. arachnoideus, H. glomeratus, H. sativus, H. tibeticus; the Mediterranean to Central Asia), Girgensohnia (5; G. bungeana, G. diptera, G. imbricata, G. minima, G. oppositiflora; southeastern European Russia to Iran and Central Asia; incl. Cyathobasis and Arthrophytum?), Cyathobasis (1; C. fruticulosa; Turkey; in Girgensohnia?), Arthrophytum (8; western and Central Asia; in Girgensohnia?), Anabasis (c 30; the Mediterranean to Central Asia); unplaced Salsoleae: Iljinia (1; I. regelii; Central Asia), Lagenantha (3; L. cycloptera, L. gillettii, L. nogalensis; northeastern Africa), Nucularia (1; N. perrinii; Algeria, Sahara), Traganopsis (1; T. glomerata; Morocco). – Caroxyloneae Akhani et Roalson in H. Akhani, G. Edwards et E. H. Roalson in Intern. J. Plant Sci. 168: 947. Jul-Aug 2007. Caroxylon (20–25; Macaronesia, the Mediterranean, northern, eastern and southern Africa, Madagascar, the Arabian Peninsula, southwestern and Central Asia, northwestern China, Xinjiang), Nanophyton (c 10; southeastern European Russia to Central Asia), Halocharis (4–7; H. clavata, H. hispida, H. sulphurea, H. violacea; southwestern and Central Asia), Kaviria (10; western Asia to Iran and Central Asia), Ofaiston (1; O. monandrum; southeastern European Russia to western Asia), Petrosimonia (9; southeastern Europe to Central Asia), Pyankovia (1; P. brachiata; southeastern Europe, Crimea and the Caucasus to Central Asia and western Siberia), Halimocnemis (6; H. karelinii, H. longifolia, H. macrantha, H. occulta, H. sclerosperma, H. villosa; southeastern European Russia to Central Asia), Piptoptera (1; P. turkestana; Central Asia), Climacoptera (40–45; southwestern Asia, Iran, Afghanistan, Central Asia); unplaced Caroxyloneae: Halarchon (1; H. vesiculosus; Afghanistan), Physandra (1; P. halimocnemis; Central Asia). – Europe, Macaronesia, the Mediterranean, Africa, southwestern and central Asia, with their highest diversity in central and southwestern Asia. Annual or perennial herbs, shrubs or trees. Leaf-succulents, halophytes. Usually C₄ metabolism (in Salsoleae usually NADP-malic enzyme subtype; in Caroxyloneae NAD-malic enzyme subtype), although in a number of clades C₃ metabolism (in S. arbusculiformis an intermediate between C₃ and C₄ metabolism). Foliar prophylls (bracteoles) present. Perianth usually with membranous wings in fruit. Perisperm poorly developed or absent. Embryo spiral. – Hammada tamariscifolia (L.) Iljin (= Salsola genistoides Juss. ex Poiret in Lam.; southern Spain) is sister to the clade [Salsoloideae+Camphorosmoideae].

Camphorosmoideae Luerss., Handb. Syst. Bot. 2: 546. Nov 1880 [‘Camphorosmeae’]

16/170–180. Chenolea (2; C. convallis, C. diffusa; Namibia, Northern, Western and Eastern Cape, KwaZulu-Natal), Spirobassia (1; S. hirsuta; Atlantic coasts of Europe to central Mediterranean, coasts of the Black Sea and the Caspian Sea), Neokochia (2; N. americana, N. californica; southwestern North America), Eokochia (1; E. saxicola; islands of Ischia, Capri and Stromboli in Italy); Grubovia (7; G. brevidentata, G. dasyphylla, G. eriophora, G. krylovii, G. melanoptera, G. mucronata, G. sedoides; Central Asia, Mongolia), ‘Maireana’ (57–60; continental and southern Australia; non-monophyletic), Dissocarpus (4; D. biflorus, D. fontinalis, D. latifolius, D. paradoxus; continental Australia), Eriochiton (1; E. sclerolaenoides; southern parts of Australia), Sclerolaena (60–65; Australia), Didymanthus (1; D. roei; southwestern Western Australia), Roycea (3; R. divaricata, R. pycnophylloides, R. spinescens; southwestern Western Australia), Eremophea (2; E. aggregata, E. spinosa; arid regions of Australia), Malacocera (4; M. albolanata, M. biflora, M. gracilis, M. tricornis; arid regions of Australia); Bassia (c 20; western Mediterranean to eastern Asia), Sedobassia (1; S. sedoides; eastern Europe to Siberia), Camphorosma (6; C. annua, C. monandra, C. monspeliaca, C. persepolitana, C. polygama, C. songorica; the Mediterranean to Central Asia). – Temperate and subtropical Eurasia, northern and southern Africa, Australia, with their highest diversity in Australia, few species in South Africa or North America. C₃ or C₄ metabolism (one species intermediate between C₃ and C₄ metabolism). Camphorosmoideae are similar to Salsoloideae, but differ through absence of floral prophylls (bracteoles), stigmas filiform and with stigmatic papillae on their entire surface, annular or folded embryo surrounding perisperm, wider pollen grains (>15 µm) with larger number of pores (usually >70) with smaller diameter (usually <2000 nm), and exine with smaller number of spinulae per operculum (>15). – Camphorosmoideae are sister to Salsoloideae.

Simplified tree of Amaranthaceae based on DNA sequence data (from numerous sources). The sister-group relationship between Polycnemoideae and the remaining Amaranthaceae is not very strong.

Maximum-likelihood tree (simplified) of Amaranthaceae based on DNA sequence data (Kadereit & al. 2006; Kadereit & Freitag 2011). [Dysphania+Teloxys] is sister-group to the Axyrideae clade in Fuentes-Bazan & al. (2012).

Nyffeler et Eggli in Taxon 59: 232. Feb 2010

Genera/species 3/c 32

Distribution Southern and southeastern Africa, Somalia, southwestern Arabian Peninsula, southwestern United States, Mexico, Bolivia, Argentina, with their largest diversity in South Africa.

Fossils Unknown.

Habit Bisexual, small shrubs or suffrutices to thick-stemmed perennial herbs. Sometimes with a basal fleshy caudex or tuberous main-root. Often stem succulents. Usually mucilaginous.

Vegetative anatomy Plants with facultative CAM (or C₄?) photosynthesis; leaves with Kranz’ anatomy. Stem epidermis usually without stomata (sometimes parallelocytic). Phellogen ab initio usually epidermal (sometimes outer-cortical). Precocious initiation of stem periderm. Phellem with lignified bands consisting of thin-walled flattened cells. Medulla often with wide-band tracheids. Cortical fibres absent. Fibre caps to stem vascular tissue absent. Secondary lateral growth normal or absent. Vessel elements with simple perforation plates; lateral pits with wide annular or helical secondary wall thickening. Imperforate tracheary xylem elements usually absent (libriform fibres present in stems of Grahamia bracteata and Talinopsis frutescens lacking wide-band tracheids). Wood rays multiseriate or absent, often with wide-band tracheids. Axial parenchyma paratracheal scanty vasicentric (in stems of Talinopsis). Thick-walled pericyclic extraxylary phloem fibre caps usually absent. Sieve tube plastids P3cf type? Nodes 1:1? Mucilaginous tracheoidal idioblasts usually frequent in wood rays (not in Grahamia). Sclereides (thick-walled) often present. Tanniniferous cells absent. Phloem parenchyma cells with phytoferritin? Parenchyma and epidermis often with numerous calciumoxalate crystals or druses.

Trichomes Hairs unicellular or multicellular, uniseriate, or absent.

Leaves Usually alternate (spiral; in Talinopsis opposite), simple, entire, often more or less succulent (especially in Grahamia), usually globose to terete (rarely flattened), with supervolute ptyxis? Stipules and leaf sheath absent. Leaves with axillary biseriate or oligoseriate hairs, bristles or a pergamentaceous scale enclosing diminutive leaves (in Anacampseros sect. Avonia). Petiole vascular bundle transection arcuate? Venation pinnate, ?-dromous. Stomata parallelocytic, usually transversely orientated. Cuticular wax crystalloids usually absent (rarely as relatively irregular platelets?). Mesophyll with mucilage idioblasts. Leaf margin entire.

Inflorescence Terminal or axillary, few-flowered thyrsoid, sometimes compact (with short internodes) or with scorpioid partial inflorescences. Involucre consisting of bracts often present. Transverse floral prophylls (bracteoles) subtending axillary flowers; median floral prophylls (in the same plane as bud-subtending prophylls) without flowers; sepaloid floral prophylls two, persistent and dry in fruit.

Flowers Actinomorphic, small to medium-sized. Hypanthium present. Hypogyny. Tepals five, with imbricate aestivation, petaloid, free or slightly connate at base (rarely absent). Nectary absent? Disc absent.

Androecium Stamens five to 15 (to c. 25). Annular androecial primordium often present in Anacampseros. Filaments free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually polypantocolpate, polyrugate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, perforate, spinulate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, ab initio multilocular, later unilocular. Stylodia (four or) five (or six). Stigmas receptive on both surfaces, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually free central (rarely basal). Ovules numerous per ovary, anatropous?, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Apical cells of megasporangium radially elongate. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus? Antipodal cells two?, ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad?

Fruit A loculicidal and/or septicidal capsule dehiscing from apex into three or six valves, with caducous exocarp usually separating from endocarp (not in Grahamia). Perianth remnants and stamens persistent and forming a dry calyptra (Grahamia, Talinopsis) or caducous (Anacampseros). Endocarp valves forming small basket.

Seeds Aril absent. Testa two-layered, sometimes winged. Exotesta in Talinaria and Anacampseros thick, with dry cells with thin unlignified walls; exotesta usually partially or entirely separating periclinally from endotesta. Endotesta? Tegmen? Perisperm starchy, usually poorly developed. Endosperm sparse or absent. Embryo usually only slightly curved, not surrounding perisperm, parallel to perisperm, well differentiated, without chlorophyll. Cotyledons two. Radicula dorsal. Germination phanerocotylar?

Cytology x = 9

DNA Intron absent from plastid gene rpl2? 6 bp deletion in plastid gene ndhF? C. 500 bp deletion in plastid gene rbcL?

Phytochemistry Betalains present. Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants.

Systematics Talinopsis (1; T. frutescens; southern United States, Mexico), Grahamia (1; G. bracteata; Bolivia, northern and central Argentina), Anacampseros (c 30; southeastern and southern Africa, Somalia, southwestern Arabian Peninsula).

Anacampserotaceae are sister-group to Cactaceae, according to Brockington & al. (2013).

The axillary appendages may by remnants of highly condensated axillary short shoots and thus homologous to the areoles in the Cactaceae (Nyffeler & Eggli 2010).

Six species of ‘Grahamia’ form a basal pectination in the analysis by Nyffeler (1997). These are now included in Anacampseros. Grahamia bracteata is sister to Anacampseros (Nyffeler & Eggli 2010).

Cladogram (simplified) of Anacampserotaceae based on DNA sequence data (Nyffeler & Eggli 2010).

Walpers in Ann. Bot. Syst. 2: 175. 15-16 Dec 1851 [‘Ancistrocladeae’], nom. cons.

Ancistrocladales Takht. ex Reveal in Novon 2: 238. 13 Oct 1992

Genera/species 1/c 20

Distribution Tropical Africa, India, Sri Lanka, eastern Himalayas to southeastern China, Southeast Asia and West Malesia.

Fossils Unknown.

Habit Bisexual, evergreen lianas with distinctly sympodial growth. Some lateral branches with circinately branched, often leaf opposite, barbs (hooks, grapnels) in one plane.

Vegetative anatomy Phellogen ab initio median-cortical. Primary cortex with thick-walled secretory cells. Cortical vascular bundles absent? Vessel elements with simple perforation plates; lateral pits alternate? Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate? (also vasicentric tracheids). Wood rays uniseriate, homocellular. Axial parenchyma apotracheal, usually in tangential bands, with fibriform vessels and spiral cells, without crystals. Wood often fluorescent. Tyloses absent. Sieve tube plastids Ss type. Nodes 3:3, trilacunar with three leaf traces. Cortical cells with pitted elongate sclereids. Wood ray parenchyma cells with silica bodies (at least in African species). Calciumoxalate crystals?

Trichomes Glandular hairs multicellular, peltate-lepidote. Vascularized glands absent.

Leaves Alternate (spiral), simple, entire, with supervolute ptyxis. Stipules very small, early caducous; leaf sheath absent. Petiole articulated. Petiole vascular bundle transection annular and with peripheral cylinder of small bundles. Venation pinnate. Stomata actinocyclocytic (surrounded by delimited subsidiary cells). Cuticular wax crystalloids as platelets. Abaxial side of lamina with small lepidote, multicellular wax glands in small pocket-like domatia. Leaf margin entire.

Inflorescence Terminal or axillary, panicle, raceme- or spike-like, racemes or spikes.

Flowers Actinomorphic to zygomorphic, small. Pedicel articulated. Half epigyny. Sepals five, with imbricate quincuncial aestivation, unequal, persistent, connate, sometimes with abaxial glands. Petals five, with contorted or imbricate aestivation, fleshy, free or connate at base. Nectaries? Disc absent.

Androecium Stamens usually ten (rarely five or 15), in one whorl (staminal primordia five?); antepetalous stamens larger. Filaments short, fleshy, connate at base, adnate to petal bases (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3(–4)-colp(or)ate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, spinulate.

Gynoecium Pistil composed of usually four (sometimes three) connate carpels. Ovary semi-inferior, unilocular. Stylodia usually four (sometimes three), free or connate at base, articulated, slightly widened in upper part. Stigmas (three or) four, crescent-shaped or pinnatifid, Dry? type. Pistillodium absent.

Ovules Placentation basal-lateral. Ovule one per ovary, hemianatropous, bitegmic, crassinucellar. Micropyle ?-stomal Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type? Endosperm development cellular. Endosperm haustoria? Embryogenesis?

Fruit A nut surrounded by hypanthium and with persistent strongly enlarged wing-like calyx lobes.

Seeds Aril absent. Seed coat exotestal? Testa membranous. Exotesta? Endotesta? Tegmen? Perisperm not developed. Endosperm hard, starchy, ruminate. Embryo small, straight, chlorophyll? Cotyledons two, strongly plicate. Germination phanerocotylar.

Cytology n = ?

DNA Intron present in plastid gene rpl2.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, delphinidin, betulinic acid, polyketide-derived naphthyl isoquinoline alkaloids (e.g., ancistrocline, dioncophylline, michellamines), and naphthoquinones (plumbagin, droserone) present. Ellagic acid?

Use Michellamines (dimerous naphthyl isoquinoline alkaloids) in Ancistrocladus korupensis are anti-HIV active substances.

Reveal et Hoogland in Bull. Mus. Natl. Hist. Nat. Paris, sér. 4, sect. B, Adansonia, 12: 205. 24 Nov 1990

Genera/species 1/8

Distribution Madagascar.

Fossils Unknown.

Habit Bisexual, evergreen trees or climbing to creeping shrubs.

Vegetative anatomy Ectomycorrhiza present. Phellogen ab initio subepidermal. Pericyclic fibres present. Primary medullary rays narrow. Continuous sclerenchyma cylinder surrounding vascular cylinder in young stems. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres (tracheids absent) with small bordered pits, non-septate. Wood rays usually uniseriate (rarely biseriate), homocellular (consisting of exclusively procumbent cells). Axial parenchyma apotracheal diffuse, or paratracheal aliform, winged-aliform, confluent or unilateral. Wood diffuse to diffuse porose, non-storied. Sieve tube plastids Ss type. Nodes 1:1, unilacunar with one leaf trace. Medullary parenchyma with sclereids. Secretory cavities? Calciumoxalate crystals and silica bodies absent.

Trichomes Hairs absent?

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole articulated. Petiole vascular bundle transection annular. Venation pinnate, brochidodromous, indistinct. Stomata paracytic? Cuticular waxes absent. Mesophyll with sclerenchymatous idioblasts containing brachysclereids. Leaf margin entire.

Inflorescence Terminal or axillary, thyrso-paniculate. Floral prophylls (bracteoles) numerous.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with imbricate aestivation, connate at base, persistent and accrescent. Petals five, with imbricate aestivation, early caducous, free, adnate to sepals. Nectary absent. Disc absent.

Androecium Stamens usually 5+5, diplostemonous (sometimes 3+3+3 or 5+5+5, triplostemonous). Filaments widened at base and connate into a ring, free from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tri- or hexacolp(oroid)ate (sometimes hexarugate, rarely dicolp[oroid]ate), shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, echinate to spinulate.

Gynoecium Pistil composed of (two or) three connate carpels. Ovary superior, usually trilocular (sometimes bilocular), with usually incomplete septa. Stylodia (two or) three, free or more or less connate. Stigmas (two or) three, punctuate, type? Pistillodium absent.

Ovules Placentation axile(-apical). Ovules usually two (sometimes four or five, rarely more than five) per carpel, anatropous, pendulous, epitropous, bitegmic, tenuinucellar? Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Nucellar cap absent. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A usually one-seeded (sometimes many-seeded) thick-walled nut, with persistent staminal filaments and strongly accrescent winged and leathery to membranous calyx, or an irregularly dehiscing capsule with weak septa.

Seeds Aril absent. Testa? Tegmen? Perisperm not developed. Endosperm almost absent. Embryo curved, well differentiated?, chlorophyll? Cotyledons two, large, spirally twisted. Germination?

Cytology n = ?

DNA

Phytochemistry Virtually unknown. Ellagic acid? Alkaloids?

Use Locally used for timber.

Nakai in J. Jap. Bot. 18: 105. 10 Mar 1942

Genera/species 1/1

Distribution Madagascar.

Fossils Unknown.

Habit Bisexual, evergreen liana. Blackening when drying.

Vegetative anatomy Phellogen superficial. Cortical fibres present. Secondary lateral growth anomalous (as concentric cylinders of vascular bundles in inner pericycle from successive cambia). Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements tracheids (and libriform fibres?) with simple pits, non-septate? Wood rays multiseriate, homocellular. Axial parenchyma paratracheal vasicentric, scanty (sometimes apotracheal, diffuse). Intraxylary phloem present. Sieve tube plastids P3cf type, with a polygonal protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Sclereids present in primary cortex. Calciumoxalate sphaeroits, rhomboidal crystals or druses present.

Trichomes Hairs absent?

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole articulated at base. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular waxes? Leaf margin entire.

Inflorescence Axillary, racemes.

Flowers Actinomorphic. Hypogyny. Tepals five, with imbricate quincuncial aestivation, sepaloid, persistent, free. Nectariferous disc annular.

Androecium Stamens c. 20–25, in several (four or five?) whorls. Filaments short, free from each other and from tepals, inserted at annular disc. Anthers subbasifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–6)-colporoidate or 3(–6)-rugate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, undulate to verrucate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular. Stylodia two, linear, slightly flattened, directed outwards to upright, somewhat connate at base. Stigmas two, flattened, papillate adaxially and along margins, type? Pistillodium absent.

Ovules Placentation basal. Ovule one per carpel, campylotropous, ascending, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A one- or two-seeded lignified loculicidal capsule.

Seeds Aril partially enclosing seed. Seed coat testal? Testal cells elongate, with sinuate anticlinal walls. Exotesta? Endotesta? Tegmen? Perisperm copious and nutritious. Endosperm sparse. Embryo peripheral, curved around perisperm, well differentiated. Cotyledons two. Germination?

Cytology n = ?

DNA

Phytochemistry Virtually unknown. Betalains? Triterpenoid saponins?

Use Unknown.

Rafinesque, Fl. Tellur. 3: 44. Nov-Dec 1837 [‘Basellides’], nom. cons.

Anrederaceae J. Agardh, Theoria Syst. Plant.: 357. Apr-Sep 1858 [’Anredereae’]; Basellineae Nakai in J. Jap. Bot. 18: 108. 10 Mar 1942 [‘Baselliineae’]; Ullucaceae Nakai in J. Jap. Bot. 18: 109. 10 Mar 1942

Genera/species 4/19

Distribution Tropical East and southeastern Africa, Madagascar, Indian Ocean islands, tropical Asia to New Guinea and islands in the Pacific, southeastern United States to tropical South America.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious; in Anredera vesicaria functionally unisexual), winding or climbing perennial herbs or lianas, slightly to distinctly fleshy. Stem bases and rhizome usually swollen and tuberous; roots sometimes tuberous.

Vegetative anatomy CAM photosynthesis? Phellogen ab initio outer-cortical. Primary vascular tissue a single cylinder of vascular strands with intraxylary phloem in larger bundles, bicollateral (at least in larger bundles); vessels only present in central portions of fascicular areas. Anomalous secondary lateral growth from concentric/successive cambia sometimes present. Vessel elements with simple perforation plates; lateral pits alternate? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma often ray-adjacent. Intraxylary phloem present. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes? Pericycle sclerenchymatous. Parenchyma with mucilaginous cells. Phloem parenchyma cells with phytoferritin? Stem epidermis with numerous calciumoxalate druses (sometimes also prismatic and octaedric crystals).

Trichomes Hairs absent.

Leaves Alternate (spiral) or nearly opposite (at stem base), simple, usually entire (rarely lobed), often somewhat fleshy, in Anredera with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate or palmate. Stomata paracytic or anomocytic. Cuticular wax crystalloids absent. Mesophyll with mucilaginous idioblasts. Leaf margin usually entire (in Tournonia glandular-serrate).

Inflorescence Terminal or axillary, spike, raceme, panicle or dichasium (in Tournonia). Floral prophylls (bracteoles) 2+2, inner pair median. Sepaloid floral prophylls two, opposite, persistent, free or partially connate at base.

Flowers Actinomorphic, small. Hypanthium present. Usually hypogyny (rarely half epigyny). Tepals (four or) five (to 13), with imbricate quincuncial aestivation, petaloid, persistent, connate at base to more than half-way up. Nectariferous disc annular, extrastaminal or intrastaminal.

Androecium Stamens (four or) five (to nine), often as many as tepals, antetepalous. Filaments widened and connate at base, in lower part adnate to tepals (epitepalous). Anthers basifixed to dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores or pore-like slits). Tapetum secretory, with multinucleate cells. Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains polypantocolpate or polypantoporate (often hexacolpate), sometimes cuboid, shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, punctate or reticulate, spinulate.

Gynoecium Pistil composed of (two or) three connate carpels. Ovary usually superior (rarely semi-inferior), unilocular (sometimes trilocular at base; in young stages trilocular). Stylodia usually three, free or connate in lower part (style sometimes single, shortly lobate). Stigmas usually three, capitate or clavate (sometimes one, trilobate), type? Male flowers with pistillodium.

Ovules Placentation basal. Ovule one per ovary or carpel, campylotropous to orthoamphitropous (anatropous?), ascending, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Nucellar cap/beak present. Megagametophyte monosporous, Polygonum type. Synergids without filiform apparatus. Antipodal cells three or more. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis chenopodiad.

Fruit A thin-walled achene (utriculus), often sunken into pergamentaceous or carnose persistent perianth or surrounded by persistent winged remnants of floral prophylls.

Seeds Aril absent. Seed coat testal-tegmic? Testa membranous. Exotesta and endotesta more or less thickened. Exotegmen? Endotegmen thickened. Perisperm scarce, surrounded by embryo. Endosperm usually copious (absent?), with clusters of starch grains. Embryo annular to cochlear, well differentiated, with chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = 12, 18, 22, 24

DNA 6 bp deletion in plastid gene ndhF.

Phytochemistry Flavonol (quercetin), flavone-C-glycosides, betacyanins, and triterpene saponins present. Ellagic acid, proanthocyanidins and cyanogenic compounds not found.

Use Ornamental plants, vegetables, dyeing of food.

Systematics Anredera (12; Texas, southern Florida, Central America, the West Indies, tropical South America to Argentina), Basella (5; B. paniculata: tropical East and southeastern Africa; B. excavata, B. leandriana, B. madagascariensis: Madagascar; B. alba: tropical Asia from India to New Guinea, possibly introduced in Africa, tropical America and on oceanic islands), Tournonia (1; T. hookeriana; Colombia), Ullucus (1; U. tuberosus; the Andes).

Basellaceae may be sister-group to Didiereaceae (Ocampo & Columbus 2010; Brockington & al. 2013) or even to the Portulacaria-Ceraria clade in Didiereaceae (Arakaki & al. 2011).

Cladogram (simplified) of Basellaceae based on morphology (Eriksson 2007).

de Jussieu, Gen. Plant.: 310. 4 Aug 1789 [’Cacti’], nom. cons.

Opuntiaceae Desv. in S. Gérardin de Mirecourt et N. A. Desvaux, Dict. Rais. Bot.: 52, 385. 12-19 Apr 1817; Cactales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 238. Jan-Apr 1820 [‘Cacteae’]; Cereaceae Spreng. ex DC. et Spreng., Elem. Philos. Bot.: 142. Jul 1821 [‘Cereae’]; Opuntiopsida Endl., Gen. Plant.: 942. Nov 1839 [’Opuntiae’]; Cactopsida Brongn., Enum. Plant Mus. Paris: xxviii, 105. 12 Aug 1843 [’Cactoideae’]; Opuntiales Endl. ex Willk., Anleit. Stud. Bot. 2: 295. 19-20 Jan 1854 [‘Opuntieae’]; Nopaleaceae Schmid et Curtman, Pflanzenr. 2: 135. 10-11 Jun 1856 [‘Nopaleae’]; Cactineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 462. 15 Nov 1895 [‘Cactales’]

Genera/species 131/1.720–1.815

Distribution Mainly arid and semiarid regions of North and South America (British Columbia and Alberta south to Patagonia); also epiphytes and lianas in rain forests and other moist forests (Rhipsalis also in tropical Africa, Madagascar, the Seychelles, the Mascarenes and Sri Lanka, possibly introduced?).

Fossils Proposed fossils of Cactaceae are ambiguous.

Habit Usually bisexual (at least in Mammillaria dioica and Selenicereus innesii functionally dioecious), usually perennial herbs (sometimes climbing or epiphytic; rarely deciduous shrubs or small trees [Rhodocactus, Pereskia]). Usually xerophytic. Almost all species are stem succulents with elongate and branched or unbranched, or almost globular pachycaul photosynthesizing stem; some genera with flattened almost foliaceous stem segments, phylloclades. Stem and branch surfaces usually with areolae – modified axillary short shoots, brachyblasts – with numerous spines – modified leaves or foliar lobes.

Vegetative anatomy CAM physiology dominating (also C₄ physiology present). Roots usually fibrous or tuberous (Peniocereus and Pterocactus with napiform nutritious roots). Stem epidermis with thick cuticle and thick cell walls, usually with parallelocytic stomata. Apical meristem very wide (often c. 0,5–1,5 mm). Phellogen ab initio usually in epidermis (rarely outer-cortical). Precocious or delayed initiation of stem periderm. Hypodermis present. Cortex and medulla with or without vascular bundles and usually modified into water-storing tissue with vacuolized parenchyma cells. Apical meristem 400–1.500 μm wide. Wide-band tracheid cells usually present in xylem strands and medulla in particular. Primary vascular tissue a cylinder of vascular bundles. Secondary lateral growth normal (inner phloem absent; secondary lateral growth absent in some groups). Vessel elements usually with simple (rarely reticulate) perforation plates; lateral pits alternate or pseudoscalariform. Imperforate tracheary xylem elements libriform fibres with simple pits (also vasicentric tracheids). Wood rays multiseriate?, wide and tall, starchy. Axial parenchyma usually paratracheal scanty. Thick-walled pericyclic extraxylary phloem fibre caps present. Sieve tube plastids P3cf type, with a central globular protein crystalloid and a subspherical ring of protein filaments, without starch. Nodes 1–?:1–>5, at least unilacunar with one to many leaf traces (trace often with two or several branches). Cortex (sometimes also medulla) with mucilaginous idioblasts and often sclereids. Laticifers in Mammillaria with white latex containing mucilaginous polysaccharides. Tanniniferous cells absent. Phloem parenchyma cells with phytoferritin? Idioblasts with crystals. Calciumoxalate (often very frequent) usually as prismatic crystals or druses (sometimes crystal sand, rarely raphides). Calciumoxalate as whewellite (CaC₂O₄^.H₂O) or weddellite (CaC₂O₄^.2H₂O). Cuticular wax crystalloids as platelets or rodlets.

Trichomes Hairs multicellular, uniseriate, or absent.

Leaves Usually almost absent (present as microscopical vestiges); in Rhodocactus and Pereskia usually alternate (spiral); leaves when present simple, entire and flat (in Rhodocactus and Pereskia) or terete (in Maihuenia and young plantlets of Opuntioideae); in Maihuenia and Opuntioideae succulent; in Pereskia with supervolute ptyxis. Stipules and leaf sheath absent. Leaves with axillary hairs (in Pereskia, Pereskiopsis and Quiabentia usually uniseriate) and sometimes multiseriate bristles or scales (hairs in Cactaceae s. str. sometimes in areolae). Petiole vascular bundles? Venation pinnate (rarely palmate) or indistinct. Stomata in Cactaceae parallelocytic, often opuntioid (subsidiary cells overlapping ends of guard cells). Cuticular wax crystalloids usually as platelets or rodlets (in Cereus clade as spiral rodlets) or as continuous crust. Mesophyll with mucilaginous idioblasts. Leaf margin entire. Extrafloral nectaries frequent at or near areolae in several genera.

Inflorescence Flowers usually solitary (rarely two or more) on lateral or subapical areolae (rarely terminal; in Pereskia cymose or racemose, panicle). Median floral prophylls absent.

Flowers Usually actinomorphic (rarely zygomorphic, e.g. Schlumbergera, Zygocactus), often large. Usually epigyny (rarely half epigyny or hypogyny). Hypanthium present or absent. Tepals few to many, spiral, with imbricate aestivation (Rhodocactus, Pereskia), usually petaloid, free or connate into perianth tube, downwards grading via sepaloid tepals and tepaloid bracts into scale-like bracts. Nectariferous disc (in, e.g., Pereskia and Rhipsalis) or nectaries at base of hypanthium and androecium (in some species of Opuntia on annular or cupular outgrowth at stylar base).

Androecium Stamens c. 20 to more than 4.000, usually at least initially spiral, later often in one or two whorls, or more than perianth tube length (staminal primordia initiated in fascicles and often continuing tepal spiral phyllotaxis). Androecial ring primordium sometimes present. Filaments free, usually adnate to perianth tube. Anthers basifixed or dorsifixed, sometimes versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia present in some genera.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, hexacolpate or polypantocolpate (in most Opuntioideae porate), usually shed as monads (rarely tetrads), tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate or reticulate, spinulate (rarely verrucate or psilate).

Gynoecium: Pistil composed of three to more than 20 connate carpels. Ovary usually inferior (in Pereskia superior or semi-inferior), usually unilocular (sometimes partially multilocular due to secondary septa). Style usually single, simple, long. Stigma trilobate to more than 20-lobate, non-papillate, Wet type. Pistillodium usually absent.

Ovules Placentation usually parietal-laminar (in Rhodocactus and Pereskia basal-laminar). Ovules numerous per ovary, usually campylotropous or circinotropous (rarely anatropous), bitegmic, crassinucellar. Micropyle endostomal. Outer integument usually approx. two (in Cereus three or four) cell layers thick. Inner integument two (or three) cell layers thick. Parietal tissue approx. one cell layer thick. Apical cells of megasporangium often radially elongate. Lateral epidermal cells anticlinally divided. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis? Polyembryony known from several genera.

Fruit Usually a berry, often dehiscent (sometimes a nut; in Copiapoa a capsule; sometimes with persistent areola, hypanthium and/or remnants of tepals). Pericarp not two-layered.

Seeds Aril (funicular) present or absent, in Opuntioideae bony. Funicle fleshy, with short apical hairs. Hilum and micropyle with various shapes (in many Cactoideae with spongy hilum/micropyle region). Testa sometimes winged. Operculum present in some representatives. Endotegmic cell walls often thickened. Perisperm copious and nutritious or almost absent. Endosperm sparse or absent. Embryo peripheral, usually strongly curved over perisperm (sometimes nearly straight or spirally twisted), well differentiated to rudimentary, without chlorophyll. Cotyledons two, usually more or less reduced. Radicula sometimes aborted. Germination phanerocotylar.

Cytology x = (9) 11 (12) – Polyploidy frequently occurring in Opuntioideae and Cactoideae. Adventitious embryony present at least in Opuntioideae.

DNA Plastid genome in at least Pereskia with inversion of 6 kb in LSC region, and c. 500 bp deletion in rbcL (cf. Portulaca!). Plastid gene infA lost/defunct (Notocactus). Intron absent from Cactoideae plastid gene rpoC1. 6 bp deletion in plastid gene ndhF.

Phytochemistry Flavonols (kaempferol, quercetin, isorhamnetin, apigenin, etc.), flavones, flavanones, dihydroflavonol glycosides, cyanidin, delphinidin, sterols, betacyanins, betaxanthins, tyramine alkaloids, phenylethylamines, pilocereine, peyote alkaloids (e.g. mescaline), and tetrahydroisoquinoline alkaloids (e.g. anhalamine, anhalidine, pellotine and anhalonidine), saponins, sapogenins and other triterpenes, and phytoferritin (in phloem parenchyma cells) present. Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants, fruits (Opuntia, Selenicereus megalanthus, Hylocereus, Stenocereus, etc.), narcotics (Lophophora williamsii), fences and hedges (e.g. Opuntia), carpentries.

Systematics Cactaceae are sister-group to Anacampserotaceae, according to Brockingty & al. (2013).

The generic and specific delimitations are notorious in Cactaceae. Hence, the species numbers within the genera are often highly provisional.

Cactaceae have undergone a tremendous increase in number of perianth parts as compared to their closest relatives (Anacampserotaceae, Portulacaceae, Talinaceae, Basellaceae, Didiereaceae and Halophytaceae). The spirally arranged perianth parts in Cactaceae may be bracteal rather than staminodial in their homology (Ronse De Craene 2007, 2008, etc.). The perianth may have evolved through differentiation and inclusion of supernumerary bracts (Ronse De Craene 2008) or through development of additional bracts. Brockington & al. (2009) suggest that this increase in perianth parts may be partially the result of a corresponding increase in meristem size and merosity of reproductive organs.

The subdivision of Opuntioideae and Cactoideae are according to Nyffeler & Eggli (2010c).

Cladogram (simplified) of Cactaceae based on DNA sequence data (Nyffeler 2007; Ogburn & Edwards 2009; Nyffeler & Eggli 2010). The clade [Opuntioideae+Maihuenioideae] collapsed in the Nyffeler & Eggli analysis.

Rhodocactus

1/5. Rhodocactus (5; R. bahiensis, R. grandifolia, R. nemorosa, R. sacharosa, R. stenantha; Mexico, the Antilles, northwestern South America, southeastern Brazil). – Phellogen outer-cortical. Precocious initiation of stem periderm. Stem epidermis without stomata or with parallel-orientated parallelocytic stomata, opuntioid except innermost cell pair, with subsidiary cells randomly arranged; leaf stomata randomly orientated; stomata opening only during rains and at night. Concentric (stratified) layers of sclereids (often thick-walled) usually present in phellem of periderm. Wide-band tracheids absent. Hypogyny or half epigyny. Pollen grains colpate. Placentation basal-laminar.

[Pereskia+[Opuntioideae+[Maihuenia+[Blossfeldia+Cactoideae]]]

Stem cuticle often thick. Epidermis persistent. Phellogen usually epidermal. Usually delayed initiation of stem periderm (not in Pereskia aculeata). Stomata usually present in stem epidermis (absent in P. nemorosa). Stem mucilage cells present. Cortical sclereids absent.

Pereskioideae Engelm. in W. H. Brewer et al., Bot. California 1: 243. 1876 [‘Peirescieae’]

1/4. Pereskia (4; P. aculeata, P. diaz-romeroana, P. horrida, P. weberiana; the Andes, Central America, northern and central South America). – Roots sometimes tuberous. Stem stomata parallel-orientated, opuntioid. Concentric (stratified) layers of sclereids (often thick-walled) usually present in phellem of periderm. Wide-band tracheids absent. Sclereids (often thick-walled) present in phloem. Lamina with supervolute ptyxis. Stomata randomly orientated, opening only during rains and at night. Hypogyny, half epigyny or epigyny. Stamens with centrifugal development from five primordia. Pollen grains polycolpate. Placentation basal-laminar.

[Opuntioideae+[Maihuenia+[Blossfeldia+Cactoideae]]]

Stem succulents with very short internodes. CAM or facultative CAM photosynthesis present. Hypodermis collenchymatous. Cortical chlorenchyma forming mesophyllar tissue with intercellular spaces. Wide-band tracheids in secondary xylem at least of seedlings with annular thickenings. Flowers solitary, axillary. Hypanthium present. Epigyny. Stamens developing from annular primordium. Placentation parietal-laminar.

Opuntioideae Burnett, Outlines Bot.: 742, 1130. Feb 1835 [’Opuntidae’]

18/170–200. Cylindropuntieae Doweld in Sukkulenty 1(2): 25. 25 Jul 1999. Austrocylindropuntia (8–10; tropical northern South America), Cumulopuntia (19–20; the Andes), Punotia (1; P. lagopus; Peru, Bolivia?), Maihueniopsis (1; M. glomerata; Peru, Bolivia, Chile, Argentina), Quiabentia (2; Q. verticillata, Q. zehntneri; eastern Brazil, Paraguay, Bolivia, Argentina), Pereskiopsis (2; P. aquosa, P. porteri; Mexico, Central America), Micropuntia (1; M. pulchella; the Mojave Desert in southwestern United States), 'Corynopuntia' (16; southwestern United States, Mexico; polyphyletic), ‘Cylindropuntia’ (20–25; southwestern United States, Mexico, Central America; non-monophyletic), Grusonia (1; G. bradtiana; southwestern United States, northwestern Mexico); Tephrocactus (13; Argentina). – Opuntieae DC., Prodr. 3: 458. Mar (med.) 1828. Miqueliopuntia (1; M. miquelii; coast of Chile), Tunilla (12; Peru, Bolivia, Chile, Argentina), Brasiliopuntia (2; B. brasiliensis, B. schickendantzii; Peru, Brazil, Paraguay, eastern Bolivia, northern Argentina), Tacinga (9; Venezuela, northeastern Brazil), ‘Opuntia’ (45–65; southwestern Canada to South America, the Galápagos Islands; non-monophyletic), Consolea (10; C. corallicola, C. moniliformis, C. spinosissima; Florida Keys, the West Indies). – Unplaced Opuntioideae Pterocactus (9; Patagonia in southern and western Argentina). – Southwestern Canada to southern Patagonia. Roots often tuberous. Stem terete, usually articulated, often flattened. Stem stomata parallel-orientated, opuntioid. Calciumoxalate as whewellite (CaC₂O₄^.H₂O). Leaves usually small, terete, succulent, caducous, present only on young stems (in Pereskiopsis large, persistent, bifacial, with lamina and petiole; in Quiabentia terete, persistent, unifacial). Areolae with glochidia (minute acicular spines with retrorse barbs). Leaf stomata parallel-oriented. Subsidiary cells of stomata not or only slightly overlapping ends of guard cells. Short hypanthium sometimes present. Pollen grains usually polyporate. Seeds covered by bony funicular aril. Outer exotestal cells with little thickened periclinal walls. Cotyledons sometimes nutrient storing organs. Deletion of plastid gene accD. – Cumulopuntia is nested in Maihueniopsis and Puna sister to this clade in the analyses by Griffith & Porter (2009).

[Maihuenia+[Blossfeldia+Cactoideae]]

Testa with interstitial craters or pits. Outer exotestal cells with strongly thickened periclinal walls.

Maihuenioideae P. Fearn, Rev. Orig. Cactus Fam.: unpaged. [53]. 10 Mar 1996

1/2. Maihuenia (2; M. patagonica, M. poeppigii; the Patagonian Andes Patagonia in southern Chile and southern Argentina). Caespitose shrubs. Phellogen epidermal. Phellogen and cork formation not delayed. epidermis persistent. Stem usually without stomata. Photosynthetic parenchyma present at base of areolar crypts. Stomata sometimes present in areolar crypts, parallelocytic. Medulla and cortex with large mucilage reservoirs. Leaves terete, succulent, persistent. Leaf with cylindrical reticulum of vascular bundles; external xylem surrounding central mucilage reservoir. Leaf stomata transversely orientated. Pollen grains tricolpate. Funicles in fruit long, mucilaginous.

[Blossfeldia+Cactoideae]

Stem stomata parallelocytic, usually unoriented (in epiphytes transverse). Leaves minute or absent. Pollen grains tricolpate to polycolpate (sometimes porate). Seeds with conspicuous spongy hilum-miropyle region. Intron absent from plastid gene rpoC1.

Blossfeldioideae Crozier in Phytologia 86: 55. 24 Sep 2004

1/1. Blossfeldia (1; B. liliputana; the eastern Andes in Bolivia and Argentina). Spines absent. Stem cuticle and epidermal wall thin, early replaced by phellogen. Photosynthetic parenchyma present at base of areolar crypts. Stem stomata few, present at base of areolar crypts. Hypodermis absent. Vascular bundles without cap of phloem fibres. Leaf stomata absent. Seeds round, with strophiolate funicular aril. Testa with a single short narrow hair per cell. – Blossfeldia is sister to Cactoideae.

Cactoideae Eaton, Bot. Dict., ed. 4: 43. Apr-Mai 1836 [‘Cacteae’]

109/1.560–1.625. Southern Canada to the southwestern United States and southwards (a few species of Rhipsalis possibly introduced by man into Africa and Madagascar). Some species are epiphytic. Primary root at least in some Cactoideae with limited growth. Stem usually ribbed (sometimes tuberculate). Stomata usually unoriented. Wide-band tracheids sometimes absent. Cortex wide, succulent. Cortical vascular bundles usually present. Cuticular wax crystalloids in Cereae as spiral-shaped rodlets. Calciumoxalate often as weddellite (CaC₂O₄^.2H₂O). Leaves absent. Flowers usually actinomorphic (sometimes zygomorphic). Hypanthium usually present. Pollen grains tricolpate to polycolpate (sometimes porate). Seeds usually non-arillate. Testa with interstitial craters or pits. Outer exotestal cells with strongly thickened periclinal walls. Hypocotyl sometimes nutrient storing organ. Intron absent from plastid gene rpoC1. Alkaloids sometimes present.

Nyffeler & Eggli (2009) found the following topology in Cactoideae: [Cacteae+[Phyllocacteae+[Rhipsalideae+[Notocacteae+Cereeae]]]].

Cacteae Rchb., Fl. Germ. Excurs. 2(2): 561. 1832

26/430–440. Geohintonia (1; G. mexicana; Nuevo Léon in northeastern Mexico; in Aztekium?), Aztekium (3; A. hintonii, A. ritteri, A. valdesii; Nuevo Léon in northeastern Mexico, incl. Geohintonia?), Echinocactus (15; southwestern United States, Mexico), Astrophytum (6; A. asterias, A. capricorne, A. caput-medusae, A. coahuilense, A. myristigma, A. ornatum; the Chihuahuan desert in southern Texas and northern Mexico); Sclerocactus (15–19; southwestern United States), Pediocactus (9; western United States), Echinomastus (6–9; E. erectocentrus, E. intertextus, E. johnsonii, E. mariosensis, E. unguispinus, E. warnockii; southwestern United States, northern Mexico); ‘Ferocactus’ (23; southwestern United States, Mexico; paraphyletic), Leuchtenbergia (1; L. principis; the Chihuahuan desert in northern Mexico; in Ferocactus?), Stenocactus (10; the Chihuahuan desert in Mexico; in Ferocactus?), Thelocactus (11; Texas, northern and central Mexico; in Ferocactus?); Epithelantha (2; E. bokei, E. micromeris; the Chihuahuan desert in southern Texas, southern Arizona and northern Mexico); Strombocactus (1; S. disciformis; eastern central Mexico), Ariocarpus (6; A. agavoides, A. bravoanus, A. fissuratus, A. kotschoubeyanus, A. retusus, A. scaphirostris; the Chihuahuan desert in southern Texas and northern Mexico), ‘Turbinicarpus’ (c 17; northern central Mexico; paraphyletic); Obregonia (1; O. denegrii; northestern Mexico), Lophophora (8; southern Texas, northern and eastern Mexico), Acharagma (3; A. aguirreanum, A. huasteca, A. roseanum; Coahuila and Nuevo Léon in northern Mexico), Rapicactus (6; Mexico); Neolloydia (5; N. conoidea, N. mandragora, N. matehualensis, N. subterranea, N. texensis; the Chihuahuan desert in Texas and northern Mexico), Cumarinia (1; C. odorata; Mexico), ‘Escobaria’ (23; southwestern Canada to Mexico, Cuba; diphyletic), Pelecyphora (2; P. aselliformis, P. strobiliformis; northern central to northeastern Mexico), Ortegocactus (1; O. macdougallii; Oaxaca province in southeastern Mexico), ‘Coryphantha’ (c 55; southwestern and southern United States, Mexico, with their highest diversity in southern United States and northern Mexico; polyphyletic), Mammillaria (>200; southwestern United States and Mexico to Colombia and Venezuela, the West Indies, with their highest diversity in Mexico). – North America incl. Mexico, with their highest diversity in the Chihuahuan desert in southern Texas and northern Mexico.

[Lymanbensonieae+[Frailea+[Phyllocacteae+[Rhipsalideae+[Notocacteae+Cereeae]]]]]

Lymanbensonieae N. Korotkova et Barthlott in Willdenowia 40: 166. 9 Dec 2010

3/c 30. Copiapoa (c 25; coastal deserts in northern Chile), Calymmanthium (1; C. substerile; Cajamarca in northern Peru), Lymanbensonia (3; L. brevispina, L. incachacana, L. micrantha; southern Ecuador, Peru, Bolivia). – Southern Ecuador to Bolivia and northern Chile. – Lymanbensonieae are sister-group to the clade [Frailea+[Phyllocacteae+[Rhipsalideae+[Notocacteae+Cereeae]]]], according to Korotkova & al. (2010).

[Frailea+[Phyllocacteae+[Rhipsalideae+[Notocacteae+Cereeae]]]]

Frailea (26; central South America).

[Phyllocacteae+[Rhipsalideae+[Notocacteae+Cereeae]]]

Phyllocacteae (Salm-Dyck) Rchb., Deutsch. Bot. Herb.-Buch: 161. Jul 1841

32/335–355. ‘Eulychnia’ (5; E. acida, E. breviflora, E. castanea, E. iquiquensis, E. procumbens; coastal deserts in Peru and northern Chile; paraphyletic), Austrocactus (5; A. bertinii, A. coxii, A. patagonicus, A. philippii, A. spiniflorus; Chile, southern Argentina), Corryocactus (c 35; southern Peru, Bolivia, northern Chile), Castellanosia (1; C. caineana; northeastern Paraguay, Cochabamba and Santa Cruz in Bolivia), Armatocereus (6; A. cartwrightianus, A. godingianus, A. laetus, A. matucanensis, A. procerus, A. rauhii; Colombia, Ecuador, Peru), Leptocereus (15; Cuba, Hispaniola, Puerto Rico), Neoraimondia (4; N. arequipensis, N. herzogiana, N. gigantea, N. macrostibas; Peru, Bolivia, northern Chile), Acanthocereus (6; A. basaniensis, A. colombianus, A. horridus, A. occidentalis, A. subinermis, A. tetragonus; Florida, southern Mexico, Central America to Colombia and Venezuela, the West Indies), Lemaireocereus (35–40; Mexico to northern South America), Bergerocactus (1; B. emoryi; coastal areas of San Diego in California and Baja California), Myrtillocactus (4; M. cochai, M. eichlamii, M. geometrizans, M. schenckii; Mexico, Guatemala), Carnegiea (1; C. gigantea; Sonoran desert in southern Arizona, southeasternmost California and northern Mexico), Pachycereus (9–13; southwestern United States, Mexico, northern Central America), Cephalocereus (14; eastern and southern Mexico), Peniocereus (19; southwestern North America, Mexico, Central America), Morangaya (1; M. pensilis; southern Baja California Peninsula in northwestern Mexico), Escontria (1; E. chiotilla; southern Mexico), Stenocereus (24; southern United States, Mexico, Central America, the West Indies, northern South America), Polaskia (2; P. chende, P. chichipe; southern Mexico), Echinocereus (55–60; South Dakota to Mexico). – Unplaced Phyllocacteae Brachycereus (1; B. nesioticus; the Galápagos Islands), Carnegiea (1; C. gigantea; Sonoran desert in southern Arizona, southeasternmost California and northern Mexico), Dendrocereus (2; Cuba: D. nudiflorus; Hispaniola: D. undulosus), Isolatocereus (1; I. dumortieri; central to southwestern Mexico), Jasminocereus (1; J. thouarsii; the Galápagos Islands), Pseudoacanthocereus (2; P. brasiliensis: Minas Gerais in Brazil; P. sicariguensis: Lara in Venezuela).– Hylocereeae Buxb. in Madroño 14: 179. 2 Mai 1958 [‘Hylocereae’]. Epiphyllum (15–20; Mexico, Central America, the West Indies, northern South America), Pseudorhipsalis (8; Central America, northern South America), Disocactus (13; southern Mexico, Central America, the West Indies, Colombia, Venezuela), Hylocereus (11–13; southern Mexico, Central America, northern tropical South America), Selenicereus (28; southern United States and Mexico, Central America, the West Indies, tropical South America), Strophocactus (3; S. chontalensis, S. testudo, S. wittii; tropical America; in Selenicereus?), Weberocereus (9; southern Mexico, Central America to Ecuador). – Southern United States to southern South America, the West Indies, the Galápagos Islands.

[Rhipsalideae+[Notocacteae+Cereeae]]

Rhipsalideae DC., Prodr. 3: 475. Mar (med.) 1828

4/c 70. Schlumbergera (9; mountains in southeastern Brazil), Lepismium (16–17; L. cruciforme, L. houlletianum, L. lorentzianum, L. lumbricoides, L. warmingianum; eastern Bolivia, southwestern Brazil, northwestern Argentina), Hatiora (7; H. cylindrica, H. epiphylloides, H. gaertneri, H. herminiae, H. pentaptera, H. rosea, H. salicornioides; eastern and southeastern Brazil), Rhipsalis (c 40; Central America, the West Indies, tropical and subtropical South America, especially Brazil, one species also in Africa, Madagascar and Sri Lanka). – Central and South America, Africa to Sri Lanka.

[Notocacteae+Cereeae]

Notocacteae Buxb. in Madroño 14: 191. 2 Mai 1958

4/125–130. Eriosyce (42; Peru, Chile), Neowerdermannia (2; N. chilensis, N. vorwerkii; Peru, Bolivia, Chile, Argentina), Parodia (80–85; eastern South America), Yavia (1; Y. cryptocarpa; Jujuy in Argentina). – South America.

Cereeae Salm-Dyck in Allg. Gartenzeitung 8: 58. 22 Feb 1840 [‘Cereastrae’]

39/545–575. Coleocephalocereus (9; eastern and southeastern Brazil), Browningia (13; the Andes in Peru, Bolivia and northern Chile), Stetsonia (1; S. coryne; Paraguay, Bolivia, Argentina), Uebelmannia (5; U. buiningi, U. gummifera, U. horrida, U. meninensis, U. pectinifera; mountains in eastern Brazil), Cereus (21; the West Indies, eastern South America), Micranthocereus (11; central and eastern Brazil), Cleistocactus (c 40; central Peru to Bolivia and northern Argentina, Paraguay, Uruguay), Yungasocereus (1; Y. inquisivensis; La Paz in Bolivia), Samaipaticereus (1; S. corroanus; Santa Cruz in Bolivia), Matucana (15; Peru), Haageocereus (c 30; deserts in Peru and northern Chile), ‘Echinopsis’ (c 130; South America; polyphyletic), Acanthocalycium (5; A. ferrarii, A. glaucum, A. klimpelianum, A. spiniflorum, A. thionanthum; Argentina; in Echinopsis?), Gymnocalycium (c 60; southeastern South America), Harrisia (18; Florida, the West Indies, northern and eastern South America to Argentina), Oreocereus (6; O. celsianus, O. doelzianus, O. hempelianus, O. pseudofossulatus, O. leucotrichus, O. trollii; Peru, Bolivia, Chile, Argentina), Rauhocereus (1; R. riosaniensis; northern Peru). – Unplaced Cereeae Arrojadoa (5; A. dinae, A. heimenii, A. marylaniae, A. penicillata, A. rhodantha; Bahia, Minas Gerais and Piaui in eastern Brazil), Arthrocereus (4; A. glaziovii, A. melanurus, A. rondonianus, A. spinosissimus; Minas Gerais and Mato Grosso in Brazil), Brasilicereus (2; B. markgrafii, B. phaeacanthus; Bahia and Minas Gerais in eastern Brazil), Cipocereus (6; C. bradei, C. crassisepalus, C. laniflorus, C. minensis, C. pleurocarpus, C. pusilliflorus; Minas Gerais in Brazil), Denmoza (1; D. rhodacantha; western and northwestern Argentina), Discocactus (11; Brazil, Bolivia), Espostoa (12; Ecuador, Peru, Bolivia), Espostoopsis (1; E. dybowskii; northern Bahia in eastern Brazil), Facheiroa (4; F. cephaliomelana, F. pubiflora, F. squamosa, F. ulei; Bahia in northeastern Brazil), Lasiocereus (2; L. fulvus, L. rupicola; Peru)?, Leocereus (1; L. bahiensis; Bahia and Minas Gerais in Brazil), Melocactus (c 35; western Mexico, Central America, the West Indies, tropical South America to southern Peru and eastern Brazil), Mila (3; M. caespitosa, M. colorea, M. nealeana; central Peru), Oroya (7; O. baumannii, O. borchersii, O. gibbosa, O. laxiareolata, O. neoperuviana, O. peruviana, O. subocculta; Peru), Pierrebraunia (2; P. bahiensis, P. brauniorum; Minas Gerais in eastern Brazil; intergeneric hybrid incl. Pilosocereus?)?, Pilosocereus (c 55; southern United States and Mexico, Central America, the West Indies, tropical South America), Praecereus (2; P. euchlorus, P. saxicola; the West Indies, South America), Pygmaeocereus (3; P. bieblii, P. bylesianus, P. familiaris; Peru), Rebutia (12–41; southern Bolivia, northwestern Argentina), Cintia (1; C. knizei; Bolivia; in Rebutia?), Stephanocereus (2; S. leucostele, S. luetzelburgii; Bahia in eastern Brazil), Weberbauerocereus (9; Peru, northern Chile). – Florida and Mexico to South America.

Phylogeny of Cactaceae based on DNA sequence data (Nyffeler 2002).

de Jussieu, Gen. Plant.: 299. 4 Aug 1789 [’Caryophylleae’], nom. cons.

Illecebraceae R. Br., Prodr. Fl. Nov.-Holl.: 413. 27 Mar 1810 [’Illecebrearum’ (Illecebreae)], nom. cons.; Paronychiaceae Juss. in Mém. Mus. Natl. Hist. Nat. 2: 386. 1815 [’Paronychieae’]; Cerastiaceae Vest, Anleit. Stud. Bot.: 271, 292. 1818 [’Cerastoideae’]; Dianthaceae Vest, Anleit. Stud. Bot.: 271, 293. 1818 [’Dianthoideae’]; Herniariaceae Martinov, Tekhno-Bot. Slovar: 307. 3 Aug 1820 [’Herniariae’]; Illecebrales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 239. Jan-Apr 1820 [‘Illecebreae’]; Ortegaceae Martinov, Tekhno-Bot. Slovar: 443. 3 Aug 1820 [’Ortegiae’]; Saginaceae Bercht. et J. Presl, Přir. Rostlin: 239. Jan-Apr 1820 [’Sagineae’]; Scleranthales Bercht. et J. Presl, Přir. Rostlin: 240. Jan-Apr 1820 [‘Sclerantheae’]; Stellariaceae Bercht. et J. Presl, Přir. Rostlin: 239. Jan-Apr 1820 [’Stellariae’]; Stellariales Bercht. et J. Presl, Přir. Rostlin: 239. Jan-Apr 1820 [‘Stellariae’]; Telephiaceae Martinov, Tekhno-Bot. Slovar: 633. 3 Aug 1820 [’Thelephides ili Telephides’]; Scleranthaceae J. Presl et C. Presl, Delic. Prag.: 66. Jul 1822 [’Sclerantheae’]; Alsinaceae Bartl. in F. G. Bartling et H. L. Wendland, Beitr. Bot. 2: 159. Dec 1825 [’Alsineae’], nom. cons.; Caryophyllopsida Bartl. in Bartl. et Wendl. in Beitr. Bot.: 137. Dec 1825 [’Caryophyllinae’]; Silenaceae (DC.) Bartl. in F. G. Bartling et H. L. Wendland, Beitr. Bot. 2: 160. Dec 1825 [’Sileneae’]; Spergulaceae Bartl. in F. G. Bartling et H. L. Wendland, Beitr. Bot. 2: 158. Dec 1825 [’Sperguleae’]; Corrigiolaceae (Dumort.) Dumort., Anal. Fam. Plant.: 44, 49. 1829; Paronychiales Link, Handbuch 2: 420. 4-11 Jul 1829 [‘Paronychiaceae’]; Telephiales Link, Handbuch 2: 45. 4-11 Jul 1829 [‘Telephiaceae’]; Silenales Lindl., Nix. Plant.: 14. 17 Sep 1833; Dianthales Burnett, Outl. Bot.: 1117. Jun 1835 [‘Dianthinae’], nom. illeg.; Minuartiaceae Mart., Consp. Regn. Veg.: 49. Sep-Oct 1835 [’Minuartieae’]; Polycarpaeaceae Mart., Consp. Regn. Veg.: 49. Sep-Oct 1835 [’Polycarpaeae’]; Lychnidaceae Döll, Rhein. Fl.: 638. 24-27 Mai 1843 [’Lychnideae’]; Sabulinaceae Döll, Rhein. Fl.: 623. 24-27 Mai 1843 [’Sabulineae’]; Caryophyllineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 461. 15 Nov 1895 [‘Caryophyllales‘]

Genera/species 82–84/2.515–2.660

Images Caryophyllaceae

Distribution Cosmopolitan although mainly temperate regions in the Northern Hemisphere, the Arctic, temperate parts of the Southern Hemisphere (including the Antarctic continent), tropical mountains, with their largest diversity in the Mediterranean and West and Central Asia.

Fossils The fossilized inflorescence of Caryophylloflora paleogenica from Tasmania has been assigned to Caryophyllaceae. Fossil pollen grains, Caryophyllidites polyoratus, are described from the Oligocene of New Zealand, and similar pollen fossils have been found in the Miocene onwards of Europe.

Habit Usually bisexual (rarely monoecious, andromonoecious, dioecious or gynodioecious), usually perennial, biennial or annual herbs (sometimes suffrutices; rarely shrubs, small trees or lianas).

Vegetative anatomy Mycorrhiza usually absent. Phellogen usually pericyclic (rarely subepidermal or in outer cortex). Pericyclic fibres present. Secondary lateral growth usually anomalous (also in roots; sometimes from concentric/successive cambia or from an inner cylidner of vascular bundles) or absent (sometimes from normal cambium). Vessel elements with simple perforation plates; lateral pits alternate or pseudoscalariform. Imperforate tracheary xylem elements usually fibre tracheids or libriform fibres (in Gymnocarpos tracheids) with usually simple (in Gymnocarpos bordered) pits. Wood rays usually absent (sometimes uniseriate or multiseriate, homocellular). Axial parenchyma usually absent (sometimes as marginal bands). Wood occasionally storied. Wood with idioblasts containing sphaerites. Intraxylary phloem present in some species. Sieve tube plastids P3c’f type, with a single polygonal central protein crystal and a subperipheral dense ring of protein filaments. Endodermis often significant. Nodes usually 1:1, unilacunar with one leaf trace, often swollen (with geotropic adjustment). Calciumoxalate as druses or crystal sand.

Trichomes Hairs unicellular or multicellular, usually uniseriate (rarely branched, dendritic); glandular hairs present or absent.

Leaves Usually opposite (rarely verticillate or alternate), simple, entire, sometimes succulent, with conduplicate or flat ptyxis. Stipules usually absent (sometimes small, membranous); leaf sheath absent. Leaf bases often connate around stem. Petiole vascular bundle transection annular, arcuate etc. Venation pinnate. Stomata usually diacytic or anomocytic (rarely anisocytic). Cuticular wax crystalloids as rodlets or platelets or as tubuli dominated by β-diketones. Leaf margin entire. Extrafloral nectaries present or absent.

Inflorescence Usually terminal or axillary, cymose combinations of dichasia and cincinni, thyrsoid (sometimes paniculate or capitate; flowers sometimes single).

Flowers Actinomorphic. Epicalyx consisting of one or more pairs of floral prophylls (bracteoles) sometimes present. Anthophore (prolonged internode between sepal and ovary, and between petals and stamens) sometimes present. Usually hypogyny (rarely perigyny). Sepals (three to) five (to 23), usually with imbricate (rarely valvate) aestivation, free or more or less connate. Petals (petaloid staminodia?) usually five (sometimes four), with usually contorted (rarely imbricate) aestivation, free, often bifid and/or clawed (sometimes fimbriate), often with a scale-like appendage, ‘corona’, in transition zone between claw and limb (‘petals’ sometimes absent); ‘corona’(in, e.g., Silene) arising from two bulges on adaxial side of ’petals’ (possibly former staminal thecae). Nectariferous disc at staminal bases or in tube formed by filament bases and petal bases or on inner side of a cupular receptacle (and sometimes nectar glands on abaxial side of base of episepalous stamens).

Androecium Stamens usually five or 5+5 (rarely one to five or more than ten), in one or two whorls, usually haplostemonous or diplostemonous (rarely 15, obdiplostemonous), usually antesepalous (rarely alternisepalous). Filaments connate at base or free from each other, often adnate to tepals (epitepalous). Anthers usually dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Outer secondary parietal cell dividing. Tapetum secretory, with binucleate or multinucleate cells. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate (rarely tricolporate) or hexa- to polypantoporate (rarely 4–12-colpate, spiraperturate or triporate), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, usually punctitegillate (sometimes anulopunctate, rarely reticulate), spinulate.

Gynoecium Pistil usually composed of two to five (to ten) connate antepetalous or antesepalous carpels, sometimes with gynophores; when three carpels then odd carpel adaxial. Ovary usually superior (rarely semi-inferior), unilocular (below and/or as young sometimes bi- to quinquelocular). Stylodia two to five, usually filiform, usually free (sometimes connate in lower part). Stigmatic area along entire or part of adaxial side of style or only at apex, papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation usually free central (as young often basally axile; rarely basal or on free septa in upper part; Uebelinia has a basal placenta with nearly circinotropous ovule). Ovules usually numerous (sometimes one or few) per carpel, hemianatropous to campylotropous, ascending, bitegmic, tenuinucellar or crassinucellar. Micropyle endostomal. Outer integument two (to five) cell layers thick. Inner integument usually two cell layers thick. Archespore unicellular to tricellular, only one cell developing. Parietal tissue sometimes three or four cell layers thick. Nucellar cap/beak present. Megagametophyte usually monosporous, Polygonum type (rarely Allium or Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells three, ephemeral; antipodal nuclei sometimes early degenerating. Chalazal caecum developed. Endosperm development ab initio nuclear. Endosperm haustoria chalazal or absent. Embryogenesis caryophyllad or solanad.

Fruit Usually a septicidal and/or loculicidal (denticidal) capsule or opening by valves (rarely a pyxidium, irregularly dehiscing capsule, a nut or a berry), sometimes with persistent calyx.

Seeds Aril usually absent (funicular aril present in Moehringia and Petrocoptis). Operculum absent. Seed coat exotestal. Exotesta usually hard and thick, often tanniniferous; outer exotestal cell walls with stalactite-shaped processes. Endotesta sometimes thickened. Exotegmen? Endotegmen usually without rod-shaped cell wall thickenings, often tanniniferous. Perisperm copious and with starch grains, surrounded by embryo. Endosperm sparse or absent. Embryo lateral-peripheral, curved around perisperm (sometimes straight, rarely spirally twisted), well differentiated, without chlorophyll. Cotyledons two. Radicula usually dorsal. Germination phanerocotylar.

Cytology x = 5–19 – Polyploidy often occurring. Protein bodies present in nucleus. Some species of Silene have a sex determination system with X and Y chromosomes.

DNA Plastid gene infA lost/defunct (Dianthus, Stellaria). Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin), flavone-C-glycosides, anthocyanins, triterpene saponins, alkaloids, pinitol, anthraquinones, phytoferritin, sterols, phytoecdysones, and cyclopeptides present. Ferulic acid present in non-lignified cell walls. Cyanidin rare. Ellagic acid, betalains and cyanogenic compounds not found.

Use Ornamental plants, detergents for washing clothes (Acanthophyllum, Saponaria).

Systematics Caryophyllaceae are sister-group to Achatocarpaceae or to the clade [Achatocarpaceae+ Amaranthaceae].

The petals have possibly androecial origin (see, e.g., Ronse De Craene & al. 1998, Ronse De Craene 2007 and 2008). However, it is ambiguous whether the absence of petals is ancestral or whether loss of petals have occurred several times in Caryophyllaceae.

The subdivision below follows Harbaugh & al. (2010) and Greenberg & Donoghue (2011), although it is still under construction (many generic names unplaced). The descriptions of the clades are mainly according to Stephens and Harbaugh & al. (2010).

Telephioideae Beilschm. in Flora 16(Beibl. 7): 92, 111. 14 Jun 1833 [‘Telephieae’]

2/16. Corrigiola (12; nearly cosmopolitan, especially Europe, Africa and Chile), Telephium (4; T. eriglaucum, T. imperati, T. oligospermum, T. sphaerospermum; the Mediterranean, northern Africa). – Subcosmopolitan. Leaves alternate (spiral). Stipules auriculate. Sepal margins membranous. Ovary with incomplete septa. Embryogenesis solanad. Corrigiola with thick-walled triangular nutlets. Telephium with irregularly three-valved and fairly thin-walled capsules. Endotegmic cells without bar-shaped thickenings. – Telephioideae are sister-group to the remaining Caryophyllaceae.

Caryophylloideae Arn., Botany: 99. 9 Mar 1832 [‘Caryophylleae’]

Leaves opposite.

Paronychieae Dumort., Fl. Belg.: 86. 1827

6/110–160. Herniaria (30–40; Europe, the Canary Islands, the Mediterranean, North Africa, Somalia, southern Africa, southwestern Asia to India, Bolivia, northern Argentina), ‘Paronychia’ (70–>110; nearly cosmopolitan, especially the Mediterranean, Turkey, southeastern United States and the central Andes in Peru to Bolivia; paraphyletic), Chaetonychia (1; C. cymosa; western Mediterranean; in Paronychia?), ‘Polycarpaea’ corymbosa (tropical and subtropical weed), Gymnocarpos (10; the Canary Islands to East Asia, northeastern Africa), Cometes (2; C. abyssinica, C. surattensis; northeastern Africa and Ethiopia to northwestern India)? – Subcosmopolitan. Stipules paired, subadaxial to petiole; paired, connate, adaxial; single, concave, adaxial or interpetiolar. Tepals hooded, with subapical abaxial awn, sometimes with membranous margins. Petals usually absent (sometimes filiform). Staminodia usually absent. Embryogenesis solanad. Fruit a nutlet. – Paronychieae are sister-group to the remaining Caryophylloideae.

[Polycarpaeae+[Sperguleae+[Rhodalsine+[[Sclerantheae+Sagineae]+[[Arenarieae+Alsineae]+[Caryophylleae+[Sileneae+Eremogoneae]]]]]]]

Corolla present.

Polycarpaeae DC., Prodr. 3: 373. Mar (med.) 1828

20/155–165. Cardionema (6; C. andina, C. burkartii, C. camphorosmoides, C. congesta, C. kurtzii, C. ramosissimum; southwestern Canada and western United States to Chile), Dicheranthus (1; D. plocamoides; the Canary Islands), Pteranthus (1; P. dichotomus; North Africa and Cyprus to Iran), Illecebrum (1; I. verticillatum; Europe, the Canary Islands, the Mediterranean), Loeflingia (4; L. baetica: southern Iberian Peninsula; L. hispanica: western Mediterranean; L. tavaresiana: southern Portugal; L. squarrosa: United States, Mexico), ’Polycarpon’ (9; warmer regions of both hemispheres; polyphyletic), ‘Polycarpaea’ (40–50; warmer regions of both hemispheres; polyphyletic),‘Scopulophila’ (2; S. parryi: Mexico; S. rixfordii: California, Nevada; diphyletic), Sphaerocoma (1; S. hookeri; northeastern Sudan, southern Arabian Peninsula, Iran), Pollichia (1; P. campestris; northern Arabian Peninsula, Ethiopia, tropical East to southern Africa), Achyronychia (1; A. cooperi; southwestern United States, Mexico), ‘Drymaria’ (c 55; western United States to Patagonia; the Galapagos Islands, one species, D. cordata, pantropical; polyphyletic; incl. Cerdia, Ortegia and Pycnophyllum?), Cerdia (1; C. virescens; Mexico; in Drymaria?), Ortegia (1; O. hispanica; the Iberian Peninsula; in Drymaria?), Pycnophyllum (c 25; the Andes; in Drymaria?), Krauseola (2; K. gillettii: northern Kenya, southern Ethiopia; K. mosambicina: Mozambique)?, Microphyes (2; M. litoralis, M. minima; Chile)?, Pirinia (1; P. koenigii; southwestern Bulgaria)?, Polytepalum (1; P. angolense; Angola)?, Stipulicida (1; S. setacea; southeastern United States, Cuba)? – Subcosmopolitan, with their highest diversity in drier subtropical regions. Stipules interpetiolar, fimbriate, from common primordium. Sepals usually hooded, awned, sometimes with membranous margins. Corolla usually absent (sometimes present, deeply lobate to entire). Stylodia connate at base. Embryogenesis solanad. Fruit usually a nutlet (sometimes a capsule). – Polycarpaeae are sister to Caryophylloideae minus Paronychieae.

[Sperguleae+[Rhodalsine+[[Sclerantheae+Sagineae]+[[Arenarieae+Alsineae]+[Caryophylleae+[Sileneae+Eremogoneae]]]]]]

Wood rays absent. Hypanthium absent. Stamens ten or more. Nectary position. Placentation sometimes axile at least basally when ovary young. Fruit usually a septicidal and loculicidal capsule (sometimes a nutlet). Fruit with two or more seeds.

Sperguleae Dumort., Anal. Fam. Plant.: 49. 1829

3/c 65. Spergula (13; temperate regions in Europe Asia, the Mediterranean, northern Patagonia), Thylacospermum (1; T. caespitosum; Central Asia, the Himalayas, western China), Spergularia (c 50; cosmopolitan). – Cosmopolitan. Stipules single, interpetiolar, connate and encircling stem below leaves. Sepals with membranous margin. Embryogenesis solanad. – Thylacospermum caespitosum is sister to Spergula (Greenberg & Donoghue 2011; Dillenberger & Kadereit 2014). Spergularia manicata (San Ambrosio Island off Chile) is a small tree.

[Rhodalsine+[[Sclerantheae+Sagineae]+[[Arenarieae+Alsineae]+[Caryophylleae+[Sileneae+Eremogoneae]]]]]

Wood rays absent. Stipules absent. Hypanthium absent. Stamens ten or more. Nectary position. Placentation sometimes axile at least basally when ovary young. Fruit usually a septicidal and loculicidal capsule (sometimes a nutlet). Fruit with two or more seeds.

Rhodalsine clade

1/1. Rhodalsine (1; R. geniculata; the Mediterranean, the Canary Islands, Morocco to northern Egypt, northern Somalia). – Rhodalsine, lacking stipules, is sister to the remaining Caryophyllaceae (Dillenberger & Kadereit 2014). Rhodalsine is sister-group to [Spergula+Spergularia], according to Kool & Thulin (2017). Thylacospermum caespitosum was not included in their analysis.

[[[Sclerantheae+Sagineae]+[[Arenarieae+Alsineae]+[Caryophylleae+[Sileneae+Eremogoneae]]]]

Sclerantheae Link ex DC., Prodr. 3: 377. Mar (med.) 1828

9–10/85–100. Pseudocherleria (12; the Caucasus and adjacent regions, arctic Asia to Japan, western North America), Triplateia (1; T. moehringioides; central Mexico), ‘Stellaria’ pro parte, Wilhelmsia (1; W. physodes; arctic regions in northeastern Asia and northwestern North America), Honckenya (1; H. peploides; coasts of temperate regions on the Northern Hemisphere, southern Patagonia), Schiedea (26; the Hawaiian Islands), Mononeuria (10; eastern North America, southwestern Greenland), Scleranthus (13; temperate regions in Europe and Asia, the Mediterranean, Ethiopia, New Guinea, Australia), Cherleria (18; central and southeastern Europe, the Mediterranean, Arctic Eurasia, the Caucasus, Central Asia, western and Arctic North America), Pentastemonodiscus (1; P. monochlamydeus; Afghanistan)? – The Northern Hemisphere, Ethiopia, New Guinea, Australia, Patagonia. Hypanthium present or absent. Sepals usually without membranous margins. Petals entire or absent. Stamens one to ten. Staminodia sometimes five. Embryogenesis solanad. Fruit a capsule dehiscing by as many valves as stylodia, or a single-seeded nutlet. – Sclerantheae are sister to Sagineae.

Sagineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1609, 1621. 1846 [‘Sagusariae’]

10/210–215. Drypis (1; D. spinosa; the Mediterranean to Lebanon); Bufonia (34; the Canary Islands, the Mediterranean, the Middle East), Mcneillia (5; M. graminifolia, M. moraldoi, M. pseudosaxifraga, M. saxifraga, M. stellata; southern and southeastern Europe, northwestern Anatolia in Turkey), Minuartia (c 55; Europe, North Africa, southwestern Asia, the Caucasus, northern India), Habrosia (1; H. spinuliflora; Syria, Iraq, western Iran), Minuartiella (4; M. acuminata, M. dianthifolia, M. elmalia, M. pestalozzae; mountain regions from Anatolia in Turkey to Nakhichivan in Azerbaijan and northern Iran), Colobanthus (c 25; mountains of southeastern Australia and Tasmania, New Zealand and adjacent islands, temperate South America and the southern Andes, the Falkland Islands, the Antarctic Peninsula and adjacent islands, South Georgia, New Amsterdam, Kerguélen, one species, C. quitensis, also in tropical South America to Mexico), Sagina (21; arctic, temperate and alpine regions on the Northern Hemisphere, mountains in East Africa and New Guinea, the Himalayas, the Andes), Facchinia (7; F. cerastiifolia, F. cherlerioides, F. grignensis, F. herniarioides, F. lanceolata, F. rupestris, F. valentina; alpine areas in Central Europe), Sabulina (c 60; southern and Central Europe, the Mediterranean, Anatolia, the Caucasus, southwestern and Central Asia, North America, Chile, Argentina). – Temperate, arctic and alpine regions on both hemispheres. Hypanthium present or absent. Sepals sometimes awned, sometimes with membranous margins (in Drypis hooded). Petals usually entire (in Drypis bifid; petals rarely absent). Stamens sometimes as many as petals. Embryogenesis solanad. Fruit usually a capsule dehiscing by as many valves as stylodia (in Drypis a single-seeded nutlet). – Sagineae are sister to Sclerantheae. Drypis may be sister to the remaining Sagineae, although it is morphologically aberrant.

[[Arenarieae+Alsineae]+[Caryophylleae+[Sileneae+Eremogoneae]]]

Hypanthium absent. Sepals connate. Petals clawed, with closed venation and adaxial ligulae. Capsule often denticidal, with twice as many teeth as stylodia. Embryogenesis usually caryophyllad.

Arenarieae Kitt., Taschenb. Fl. Deutschl., ed. 2, 2: 981. 1844

2/200–>210. Moehringia (c 30; temperate Europe, Asia and North America), Arenaria (170–>180; temperate regions in Europe and Asia, mountains in East Africa, North America, Mexico, Central America). – The Northern Hemisphere, Central and western South America. Petals entire (rarely absent). Fruit a valvicidal capsule, dehiscing by twice as many valves as stylodia. Oily funicular aril present in Moehringia. Cotyledons incumbent. – Arenarieae are sister-group to Alsineae.

Alsineae Lam. et DC., Syn. Plant. Fl. Gall.: 392. 30 Jun 1806

12/400–410? Lepyrodiclis (3; L. holosteoides, L. stellarioides, L. tenera; Turkey to the Himalayas), Pseudostellaria (17; Europe, Afghanistan, Central Asia and western China to the Korean Peninsula and Japan, Canada, United States), Odontostemma (c 65; China, Tibet, Sikkim), Shivparvatia (7; S. ciliolata, S. forrestii, S. glanduligera, S. ludlowii, S. ramellata, S. rhodantha, S. stracheyi; northern India, the Himalayas, southwestern China), Holosteum (5; H. glutinosum, H. kobresietorum, H. marginatum, H. polygamum, H. umbellatum; temperate regions in Europe and Asia, Ethiopia), Moenchia (4; M. coerulea, M. erecta, M. graeca, M. mantica; West and Central Europe, the Mediterranean), Cerastium (180–190?; nearly cosmopolitan), Stellaria (c 120; Europe, the Mediterranean, mountains in Africa, Asia, few species cosmopolitan), Brachystemma (1; B. calycinum; the Himalayas, China, Indochina)?, Pseudocerastium (1; P. stellarioides; Anhui in eastern China)?, Pycnophyllopsis (4; P. cryptantha, P. keraiopetala, P. muscosa, P. tetrasticha; the Andes in Bolivia and Patagonia)?, Thurya (1; T. capitata; southwestern Asia)? – Subcosmopolitan, with their largest diversity in Europe to East Asia. Hypanthium usually absent. Sepals sometimes with membranous margins. Petals usually retuse to deeply bifid (rarely absent). Fruit usually a denticidal capsule (rarely a single-seeded nutlet). – Alsineae are sister to Arenarieae.

[Caryophylleae+[Sileneae+Eremogoneae]]

Veins at leaf apex intramarginal. Sepals connate. Anthophore (prolongation between sepals and remaining floral parts) sometimes present. Petals with contorted aestivation, clawed, often with retuse apex, with closed venation. Coronal scale present or absent. Nectary usually adaxial on stamens. – The sister-group relationships among Caryophylleae, Eremogoneae and Sileneae are not clarified.

Caryophylleae Lam. et DC., Syn. Plant. Fl. Gall.: 386. 30 Jun 1806

13/665–685. Psammosilene (1; P. tunicoides; Yunnan); Gypsophila (c 150; temperate regions of Eurasia, eastern Mediterranean, the Middle East, Egypt, Somalia, the Arabian Peninsula, one species, G. australis, in Australia and New Zealand; polyphyletic), Saponaria (c 30; temperate regions of Eurasia, with the highest diversity in the Mediterranean and the Middle East), Acanthophyllum (90–100; southwestern, southern and Central Asia, Siberia), Heterochroa (6; H. antoninae, H. desertorum, H. microphylla, H. petraea, H. turkestanica, H. violacea; Kazakhstan, western Siberia, Altai, the Russian Far East, the Kamchatka Peninsula, Mongolia, northern China), Cyathophylla (2; C. chlorifolia: mountains in Greece and Turkey; C. viscosa: Azerbaijan), Petroana (2; P. montserratii: the Iberian Peninsula; P. montana: Somalia, Yemen, Socotra, Oman), Bolanthus (c 10; southern Bulgaria, Greece, southern and central Turkey, Syria, Lebanon, Palestine), Psammophiliella (5; P. filipes, P. floribunda, P. kermanensis, P. muralis, P. picta; Europe to Central Asia), Balkana (1; B. spergulifolia; western and central Balkan Peninsula), Graecobolanthus (8; G. chelmicus, G. creutzburgii, G. fruticulosus, G. graecus, G. intermedius, G. laconicus, G. thessalus, G. thymifolius; Greece, Crete), Petrorhagia (c 30; Europe, the Canary Islands, the Mediterranean, southwestern and Central Asia to Kashmir), Dianthus (330–340; temperate regions in Europe and Asia, the Mediterranean, mountains in East Africa, one species, D. repens, also in northwestern North America). – Eurasia, Africa, one species in North America and one species in Australia and New Zealand. “Epicalyx” (consisting of floral prophylls) sometimes present. Calyx tubular. Sepals without commissural veins. Petals with usually dextrorse-contorted (sometimes imbricate) aestivation. Pistil composed of two (or three) connate carpels. n = 12–15, 16. – Psammosilene is sister to the remaining Caryophylleae (Madhani & al. 2018). Caryophylleae are possibly sister-group to [Sileneae+Eremogoneae]. On the other hand, Sadeghian & al. (2015) recovered Caryophylleae as sister to either Eremogoneae or Sileneae, although with low support.

[Sileneae+Eremogoneae]

Sileneae DC., Prodr. 1: 351. Jan (med.) 1824

2/520–530. Agrostemma (3; A. brachylobum, A. githago, A. gracile; temperate regions of Europe and Asia, the Mediterranean), Silene (520–530; Europe, Macaronesia, the Mediterranean, Africa, Asia, North America, the Hawaiian Islands). – North temperate to subtropical regions, montane regions in Africa. Calyx tubular. Sepals with commissural veins. Petals with usually contorted (sometimes imbricate) aestivation. Placentation sometimes axile at base. – Sileneae are sister-group to Eremogoneae. – Evolution of protease gene clpP often accelerated.

Eremogoneae Rabeler et W. L. Wagner in D. T. Harbaugh et al., Intern. J. Plant Sci. 171: 196. 14 Jan 2010

1–3/90–100. Eremogone (90–95; temperate regions on the Northern Hemisphere). – Western North America, Eurasia, the Middle East. Usually perennial herbs. Leaves filiform to subulate. Hypanthium usually poorly developed. Sepals free (calyx not tubular), with membranous margins. Ovary usually poorly semi-inferior. Cotyledons accumbent. – Eremogoneae are sister-group to Sileneae. Phlebanthia (here included in Eremogone) is sister to Eremogone, according to Harbaugh & al. (2010). – Dolophragma (5–7; D. globiflorum, D. juniperinum, D. oreophilum, D. polytrichoides, D. przewalskii, D. smithianum; the Himalayas, western China) may be sister to Eremogone, according to Sadeghian & al. (2015).

Unplaced Caryophyllaceae

Kabulia (1; K. akhtarii; Afghanistan).

Phylogeny of Caryophyllaceae based on DNA sequence data (Harbaugh & al. 2010; Greenberg & Donoghue 2011).

Phylogeny of Caryophyllaceae based on Dillenberger & Kadereit (2014).

Cladogram (simplified) of Caryophyllaceae based on DNA sequence data (Fior & al. 2006).

Radlkofer in Engler et Prantl, Nat. Pflanzenfam., III, 5: 462. 23 Apr 1896 [‘Didiereae’], nom. cons.

Portulacariaceae (Fenzl) Doweld, Tent. Syst. Plant. Vasc.: xlii. 23 Dec 2001

Genera/species 6/20

Distribution Southwestern Madagascar, southern and eastern Africa.

Fossils Unknown.

Habit Bisexual (Calyptrotheca, Portulacaria) or dioecious (in Ceraria and Decaryia rarely gynodioecious), evergreen or deciduous trees or shrubs (rarely climbing). Many species are cactus-like xerophytic stem succulents, often with short shoots (brachyblasts) modified into a small and definite number of spines.

Vegetative anatomy CAM or facultative CAM photosynthesis present. Stem epidermis with or without parallelocytic stomata. Phellogen ab initio superficial? Precocious initiation of periderm. Phellem in at least Ceraria and Portulacaria with lignified bands consisting of thin-walled flattened cells. Primary medullary strands wide. Secondary lateral growth normal. Vessel elements with simple perforation plates; lateral pits alternate to pseudoscalariform or scalariform, simple pits. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate (also vasicentric tracheids). Wide-band tracheids sometimes present. Wood rays multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal vasicentric or banded. Thick-walled pericyclic extraxylary phloem fibre caps present. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Intraxylary phloem absent. Nodes 1:1? Cortex and medulla often with tanniniferous cells (not in Calyptrotheca), mucilage cells and secretory mucilage ducts. Sclereids (often thin-walled, elongated with lignified walls) often present. Mucilaginous idioblasts and tanniniferous cells abundant. Phloem parenchyma cells with phytoferritin? Parenchyma and epidermis often with numerous prismatic crystals or druses of calciumoxalate.

Trichomes Stem epidermis often with papillae or multicellular, uniseriate hairs. Spines without trichomes.

Leaves Alternate (spiral) or opposite, simple, entire (caducous in some species; in Calyptrotheca succulent), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation usually pinnate (in Ceraria and Portulacaria palmate). Stomata parallelocytic (transversely orientated). Cuticular wax crystalloids as cross-bars or rodlets. Mesophyll with mucilaginous idioblasts. Leaf margin entire.

Inflorescence Terminal or axillary, dichasium, capitate, fasciculate, panicle etc., or flowers solitary axillary. Floral prophylls (bracteoles) 2+2, with inner pair (subtending flower) median. Some genera with two free, sepaloid (sometimes coloured, petaloid), often persistent floral prophylls subtending flower. Transverse floral prophylls absent. Sepaloid floral prophylls with imbricate aestivation.

Flowers Actinomorphic. Usually hypogyny (rarely half epigyny). Tepals four or five, with imbricate or decussate aestivation, petaloid, usually free (in Portulacaria connate into a tube), or absent. Nectariferous disc intrastaminal, annular.

Androecium Stamens four or five, in one whorl, alternitepalous, or seven to numerous (in Calyptrotheca up to c. 60), in several whorls; developed from annular primordium. Filaments free or connate at base into ring of adaxial nectaries, usually free from tepals (in Portulacaria adnate to sepals, episepalous). Anthers dorsifixed, often versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Female flowers sometimes with staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually polypantoporate or tetra- to heptazonocolpate (in Ceraria and Portulacaria tricolpate; in Calyptrotheca polypantoporate), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum (due to fusions?), microperforate, spinulate and globulate.

Gynoecium Pistil composed of (two or) three (or four) connate carpels (in Ceraria and Portulacaria a single carpel). Ovary usually superior (rarely semi-inferior), usually ab initio trilocular (sometimes bilocular or quadrilocular), later secondarily unilocular (median/adaxial locule) due to degeneration of remaining locules. Style single, usually simple (in Portulacaria trifid). Stigmas more or less peltate, fimbriate (bifid or) trifid (or quadrifid), type? Male flowers sometimes with pistillodium.

Ovules Placentation basal. Ovules one to six (Calyptrotheca) per ovary (when carpel adaxial) or one per carpel, campylotropous to hemicampylotropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle endostomal?, directed downwardly. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis chenopodiad.

Fruit Usually a one-seeded achene (in some groups enclosed inside two dry sepaloid bracteoles; in Ceraria a samara; dehiscence in Calyptrotheca circumscissile at base and splitting upwards into valves in upper part, with strongly accrescent calyx).

Seeds Seeds with funicular strophiole or (in, e.g., Calyptrotheca) aril. Testa? Tegmen? Perisperm sparse or absent. Endosperm entirely or almost entirely absent. Embryo large, curved around perisperm, well differentiated, chlorophyll? Cotyledons two, often fleshy. Germination phanerocotylar.

Cytology n = 8–12(–17), 22, 24, 86, 120 – Polyploidy frequently occurring.

DNA 6 bp deletion in plastid gene ndhF.

Phytochemistry C-methylated flavonoids, cyanidin, delphinidin, betacyanins, betaxanthins, and phytoferritin present. Triterpene saponins? Cyanogenic compounds not found.

Use Ornamental plants.

Systematics Portulacaria (7; P. afra, P. armiana, P. carrissoana, P. fruticulosa, P. longipedunculata, P. namaquensis, P. pygmaea; tropical and southern Africa); Calyptrotheca (2; C. somalensis, C. taitensis; northeastern tropical Africa); Alluaudiopsis (2; A. fiherensis, A. marnieriana; southern and southwestern Madagascar), Alluaudia (6; A. ascendens, A. comosa, A. dumosa, A. humbertii, A. montagnacii, A. procera; southern and southwestern Madagascar), Decarya (1; D. madagascariensis; southwestern Madagascar), Didierea (2; D. madagascariensis, D. trollii; southern and southwestern Madagascar).

Didiereaceae may be sister-group to Basellaceae (Ocampo & Columbus 2010; Brockington & al. 2013). However, some analyses (e.g. Arakaki & al. 2011) include Basellaceae in Didiereaceae.

DNA analyses give the topology [Portulacaria+[Calyptrotheca+[Alluaudia+Alluaudiopsis+Decarya+Didierea]]].

Lignified bands consisting of thin-walled flattened cells are present in the phellem in Ceraria fruticulosa and Portulacaria afra (Ogburn & Edwards 2009).

The axillary appendages may be remnants of highly condensed axillary short shoots and homologous to the areolae in Cactaceae (Nyffeler & Eggli 2010).

Cladogram of Didiereaceae based on DNA sequence data (Applequist & Wallace 2000; Nyffeler & Eggli 2010).

Airy Shaw in Kew Bull. 6(1951): 333. 26 Jan 1952, nom. cons.

Dioncophyllales Takht. ex Reveal in Phytologia 74: 174. 25 Mar 1993

Genera/species 3/3

Distribution Tropical West and Central Africa.

Fossils Unknown.

Habit Bisexual, evergreen lianas or climbing and scrambling shrubs (climbing up to c. 70 m high). Carnivorous (at least Triphyophyllum).

Vegetative anatomy Phellogen? Cortex with vascular bundles and massive fibrous bands. Secondary lateral growth in Dioncophyllum anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements tracheids and fibre tracheids with bordered pits (also vasicentric tracheids). Wood rays usually uniseriate, homocellular. Axial parenchyma apotracheal, diffuse, or paratracheal, scanty vasicentric. Intraxylary phloem present. Sieve tube plastids S type. Nodes? Crystals?

Trichomes Glandular hairs multicellular, insensitive, stalked or sessile, secreting a viscous mucilage with and without proteolytic enzymes, respectively. Glandular heads in Triphyophyllum consisting of two layers of secretory cells and inside these layers an endodermis; glandular stalk usually vascularized (xylem and phloem).

Leaves Alternate (spiral), simple, entire, often coriaceous, ’midrib’ leaves in Triphyophyllum with circinate ptyxis. Stipules and leaf sheath absent. Rosette leaves long, densely parallelodromous; higher leaves with short lamina and, in Triphyophyllum, stalked and sessile multicellular vascularized glandular hairs (with proteolytic enzymes in secretory drips) on prolonged mid-vein on abaxial side of lamina; terminal leaves (on climbing parts of shoots) with paired recurved apical barbs (hooks, grapnels). Dioncophyllum and Habropetalum with simple leaves and barbed leaves. Triphyophyllum with barbed leaves on older shoots, simple leaves on short shoots, and abaxially circinate filiform densely glandular leaves on young short shoots. Petiole vascular bundle transection crescent-arcuate. Venation usually pinnate (on some leaves of Triphyophyllum parallelodromous); mid-vein usually prolonged and dichotomizing in two recurved barb- or hook-shaped tendrils. Stomata actinocyclocytic or cyclocytic. Cuticular wax crystalloids as platelets. Leaf margin entire or crenulate.

Inflorescence Axillary or supra-axillary, cymose, of various shape.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with open or valvate aestivation, persistent, free or connate at base into a tube. Petals five, with contorted aestivation, caducous, free. Nectary absent. Disc absent.

Androecium Stamens ten or 25 to 30 (in Dioncophyllum sometimes five antepetalous). Filaments free or slightly connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective somewhat prolonged. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3- or 4-colp(or)ate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, spinulate or scabrate.

Gynoecium Pistil composed of two (Triphyophyllum) or five connate carpels; carpels sometimes opening as seed develops. Ovary superior, unilocular. Stylodia two (Triphyophyllum) or five, filiform, short, free or slightly connate at base, or absent. Stigmas two capitate (Triphyophyllum), or five punctate, capitate or plumose (Triphyophyllum), non-papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal. Ovules c. 30 to more than 100 per ovary, anatropous, bitegmic, crassinucellar. Micropyle exostomal? Outer integument ? cell layers thick, forming an envelope. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development nuclear? Megagametophyte and endosperm transversely elongate. Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule, dehiscing before maturation with two (Triphyophyllum) or five valves exposing unripe seeds.

Seeds Seeds flattened, on elongated stiff funicles, often broadly winged. Aril absent? Seed coat thick. Testa? Tegmen? Perisperm not developed. Endosperm copious, starchy, surrounding larger part of embryo. Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, semicircular. Germination cryptocotylar.

Cytology n = 12, 18? (Triphyophyllum peltatum)

DNA Intron present in plastid gene rpl2.

Phytochemistry Betulinic acid, polyketide-derived naphthyl isoquinoline alkaloids (e.g. ancistrocladine, dioncophylline, habropetaline and habropetaloic acid), cyclopentenoid cyanogenic glycosides, naphthoquinones (plumbagin), cis- and trans-isoshinanolone, and habropetalal present. Ellagic acid?

Use Medicinal plants.

Salisbury, Parad. Lond. 2: ad t. 95. 1 Feb 1808 [‘Drosereae’], nom. cons.

Droserales Bercht. et J. Presl, Přir. Rostlin: 217. Jan-Apr 1820 [‘Drosereae’]; Dionaeaceae Raf., Fl. Tellur. 3: 35. Nov-Dec 1837 [‘Dionidia’]; Aldrovandaceae Nakai in J. Jap. Bot. 24: 10. Dec 1949

Genera/species 3/probably >200

Distribution Cosmopolitan except polar and arid regions, with their largest species diversity in Australia and New Zealand.

Fossils Seeds assigned to Aldrovanda (c. fossil 20 species described) have been found in the Oligocene and the Miocene of Siberia and the Eocene of Europe. Some cenozoic pollen grains assigned to Droseraceae have been described under the name of Saxonipollis. Fossil pollen of Drosera have been reported from the Miocene of New Zealand and Europe.

Habit Bisexual, perennial or annual herbs (in Drosera sometimes climbing). Corm or tuberous rhizome present in some species of Drosera. Most species are hygrophytes; Aldrovanda is aquatic with submersed leaves. Carnivorous; Aldrovanda: ‘snaptraps’ with c. 20 trigger hairs per foliar lobe; Dionaea: ‘snaptraps’ with three trigger hairs per foliar lobe; Drosera: ‘flypaper traps’.

Vegetative anatomy Mycorrhiza absent. Main root usually ephemeral and replaced by adventitious roots (roots absent in mature plants of Aldrovanda). Phellogen? Young stem with separate vascular bundles in one or two cylinders. Medullary vascular bundles present in some species of Drosera; medullary rays wide. Cambium and secondary lateral growth absent. Vessel elements with simple perforation plates; lateral pits? Imperforate tracheary xylem elements? Wood rays absent. Axial parenchyma? Sieve tube elements S type. Nodes 1:1, unilacunar with one leaf trace. Crystals?

Trichomes Eglandular hairs present as multicellular uniseriate trigger hairs in Aldrovanda, Dionaea and Drosera glanduligera. Characteristic multicellular glandular hairs and sessile glands. Glands in Aldrovanda sessile, non-vascular, quadrifid, adaxial; in Dionaea sessile, non-vascular, adaxial and stellate, abaxial; and in Drosera stalked, vascular (with xylem, without phloem), adaxial. Tip of ‘catching glands’ in Drosera consisting of two layers with endodermis inside.

Leaves Usually alternate (usually in a basal rosette; in Aldrovanda verticillate), simple, usually entire (in Drosera sometimes once or several times dichotomously lobed), often incurved and with adaxially circinate ptyxis; leaves in Aldrovanda and some species of Drosera bifid, articulated along mid-vein. Stipules intrapetiolar or absent; leaf sheath absent. Petiole in Dionaea and some species of Drosera horizontally widened. Petiole vascular bundle transection annular, arcuate etc. Venation? Stomata usually anomocytic (sometimes tetracytic or actinocytic) or absent. Cuticular wax crystalloids as threads (round in cross-section) forming felt-like covering. Leaf margin usually entire (in Dionaea coarsely serrate). Lamina in Aldrovanda and Dionaea modified into a ’snaptrap’ with foldable laminal halves provided with sensitive trigger hairs, or margins and adaxial side of lamina provided with sensitive viscid long-stalked glandular hairs and sessile glands, secreting proteolytic enzymes; leaf margins in Drosera bending inwards when touched.

Inflorescence Usually terminal, thyrsopaniculate, scorpioid or cincinnus (flowers in Aldrovanda solitary). Bracts and floral prophylls (bracteoles) sometimes absent.

Flowers Actinomorphic. Hypogyny. Sepals (four or) five (to eight), with imbricate aestivation, marcescent, usually connate at base. Petals (four or) five (to eight), with convolute (imbricate or contorted?) aestivation, short-stalked, marcescent, free. Nectaries absent? Disc absent.

Androecium Stamens as many as sepals, antesepalous, (four or) five (Drosera, Aldrovanda) or ten to 15 (to 20) (Dionaea). Filaments usually free (in Dionaea connate at base), free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, usually extrorse (rarely introrse), longicidal (dehiscing by longitudinal slits); connective sometimes expanded. Tapetum usually secretory (sometimes amoeboid-periplasmodial). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains triporate (Aldrovanda) or 10–30-porate (Dionaea, Drosera), shed as tetrads (in Drosera usually with radial discs), usually bicellular (sometimes tricellular) at dispersal. Exine usually tectate (sometimes intectate), with columellate infratectum, usually spinulate (not in Dionaea), in Drosera often operculate.

Gynoecium Pistil composed of three (to five) paracarpous and connate carpels; median carpel abaxial. Ovary superior, unilocular. Stylodia three (to five), free, in Aldrovanda and Drosera simple or more or less bilobate (style in Dionaea single, usually simple). Stigmas three (to five), expanded, capitate, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually parietal (in Dionaea basal). Ovules one to numerous per carpel, anatropous, ascending, bitegmic, usually crassinucellar (in some species of Drosera tenuinucellar). Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megasporangial epidermal cells enlarged. Parietal tissue often absent. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Megagametophyte haustorium present. Endosperm development nuclear. Endosperm haustoria? Embryogenesis caryophyllad to solanad or onagrad to asterad.

Fruit Usually a loculicidal capsule (in Aldrovanda a nutlet).

Seeds Aril absent. Seed coat testal (Aldrovanda), exotestal (Dionaea) or endotestal (Drosera). Exotesta thick (Aldrovanda, Dionaea) or thin (Drosera), often palisade. Endotestal cell walls occasionally with U-shaped thickenings. Exotegmen crushed. Endotegmic cells small, tanniniferous, sclerotized or mucilaginous. Megasporangium in Drosera strongly prolonged during seed development. Perisperm not developed. Endosperm copious, oily (and starchy?). Embryo straight, well differentiated, sometimes short, chlorophyll? Cotyledons two; their tips with haustorial function. Germination phanerocotylar or cryptocotylar. Radicula usually ephemeral.

Cytology n = 5 or more (Drosera), n = 16 (Dionaea), n = 24 (Aldrovanda) – Polyploid and aneuploid series frequent in Drosera (n = 5–8, 10–12, 14–19); chromosomes less than 1,5 µm long (in Dionaea up to 6 µm long). Drosera and Aldrovanda (not Dionaea) have diffuse centromeres.

DNA Plastid gene infA lost/defunct (Dionaea). Intron in plastid gene rpl2 rarely lost. Mitochondrial coxI intron present (Dionaea).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), hyperoside (quercetin 3-O-galactoside), cyanidin, ellagic acid, gallic acid, proanthocyanidins (prodelphinidins), cyanogenic compounds, and naphthoquinones (plumbagin, droserone and 7-methyljuglone) present. Alkaloids and saponins not found.

Use Ornamental plants, medicinal plants.

Systematics Drosera (probably >200; cosmopolitan, especially Australia), Dionaea (1; D. muscipula; North and South Carolina), Aldrovanda (1; A. vesiculosa; central and eastern Europe, Africa, Asia to Queensland).

Droseraceae may be sister to Nepenthes (Nepenthaceae) or, possibly, to the clade [Nepenthaceae+[Drosophyllaceae+[Ancistrocladaceae+Dioncophyllaceae]]].

Dionaea is sister to Aldrovanda. The two clades share the synapomorphies leaves with ’snaptraps’, sensitive multicellular trigger hairs, sessile non-vascularized glands, and large smooth seeds with a thick exotesta. Dionaea differs from Aldrovanda by presence of stellate glands, 15 stamens, multiaperturate non-spinulate pollen grains with alternate pores at tetrad insertion, connate stylodia and basal placentation.

Cladogram of Droseraceae based on DNA sequence data (Cameron & al. 2002).

Chrtek, Slavíková et Studnička in Preslia 61: 122. 10 Apr 1989

Genera/species 1/1

Distribution Portugal, southern Spain, northern Morocco.

Fossils Unknown.

Habit Bisexual, suffrutex with long-lived taproot (sometimes a biennial herb). Carnivorous. Leaves with sweet scent.

Vegetative anatomy Mycorrhiza probably absent. Phellogen ab initio superficial. Cortical vascular bundles inverted. Secondary lateral growth normal. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits. Wood rays uniseriate or biseriate (consisting of mostly upright elements), usually homocellular. Axial parenchyma apotracheal diffuse (sometimes diffuse-in-aggregates), or paratracheal scanty vasicentric. Sieve tube plastids S type? Nodes? Crystals?

Trichomes Eglandular hairs absent. Stem and leaves covered with rows of stalked mucilage tentacles (mucilage hygroscopic) and irregularly distributed sessile digesting glands, otherwise glabrous. Glandular head composed of two layers of secretory cells and inside these endodermis; glandular stalk vascularized (with xylem and phloem).

Leaves Alternate (spiral), simple, entire, linear, with abaxially circinate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation? Stomata? Cuticular wax crystalloids? Lamina immobile, on margins and abaxial side with sessile glands, and longitudinal rows of stalked viscid non-sensitive glandular hairs secreting proteolytic enzymes, and immobile sensitive hairs for capturing insects. Leaf margin entire?

Inflorescence Terminal, few-flowered, thyrso-paniculate.

Flowers Actinomorphic, often large. Hypogyny. Sepals five, with imbricate aestivation, glandular, connate at base. Petals five, with contorted aestivation, marcescent, free. Nectaries? Disc absent.

Androecium Stamens 5+5, antesepalous longer than antepetalous. Filaments free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains polypantoporate with c. 40 pores, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, microperforate, spinulate.

Gynoecium Pistil composed of five paracarp and connate antesepalous carpels. Ovary superior, unilocular. Stylodia five, free. Stigmas capitate, papillate, Dry type? Pistillodium absent.

Ovules Placentation basal. Ovules numerous per ovary, anatropous, bitegmic, crassinucellar, with long funicle. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type? Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Septicidal and in upper part loculicidal capsule, with antesepalous valves.

Seeds Aril absent. Seeds operculate. Seed coat testal-tegmic. Testa thick. Exotesta non-palisade. Endotestal cells with crystals and U-shaped wall thickenings. Exotegmen thick-walled. Endotegmen partially sclerotized. Perisperm not developed. Endosperm copious, fleshy, starchy. Embryo small, cordate, well differentiated, chlorophyll? Cotyledons two; cotyledon tips with haustorial function. Germination cryptocotylar.

Cytology n = 6 – Chromosomes more than 15 µm long (much larger than in Droseraceae). Centromere localized.

DNA Intron present in plastid gene rpl2.

Phytochemistry Flavones (e.g. luteolin), proanthocyanidins, naphthoquinones (plumbagin etc.), and inulin present.

Use Ornamental plant.

Desvaux in S. Gérardin de Mirecourt et N. A. Desvaux, Dict. Rais. Bot.: 188. 12-19 Apr 1817 [’Frankeniae’], nom. cons.

Frankeniales Link, Handbuch 2: 229. 4-11 Jul 1829 [‘Frankeniaceae’]

Genera/species 1/70–80

Distribution Subtropical and warm-temperate dry regions.

Fossils Unknown.

Habit Usually bisexual (sometimes polygamomonoecious), evergreen shrubs, suffrutices or perennial (sometimes annual) herbs. Usually halophytic or xerophytic, sometimes gypsophilous or calciphilous.

Vegetative anatomy Mycorrhiza usually absent. Phellogen subepidermal or pericyclic (intraxylary cork-tissue present in some species). Secondary lateral growth in some species of Frankenia anomalous (from concentric/successive cambia). Fibriform vessel elements present. Vessel elements with simple perforation plates; lateral pits alternate, simple pits? Imperforate tracheary xylem elements libriform fibres usually with simple pits (fibres in some species replaced by axial parenchyma; also vasicentric tracheids). Wood rays absent. Axial parenchyma? Wood elements partly storied. Sieve tube plastids S type. Nodes? Crystals?

Trichomes Hairs usually unicellular, uniseriate, sometimes stellate as tufts; salt glands present.

Leaves Opposite, simple, entire, linear, often ericoid, with ? ptyxis. Stipules absent; leaves in each pair fused by a common sheath. Petiole vascular bundles? Leaf single-veined. Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids absent. Mesophyll often with sclerenchymatous idioblasts in association with vascular strands. Epidermis often with salt-secreting glands (consisting of six crescent cells in two layers of three). Leaf margin entire, usually reflexed.

Inflorescence Terminal or axillary, cymose of various shapes, or flowers solitary axillary.

Flowers Actinomorphic, small. Hypogyny. Sepals usually five (sometimes four, six or seven), with induplicate-valvate aestivation, persistent, usually connate into a tube (calyx in Frankenia triandra campanulate or urceolate). Petals usually five (sometimes four, six or seven), with imbricate aestivation, clawed, usually with adaxial scale-like appendage, ligule, with nectary on stipe, free. Disc absent.

Androecium Stamens (three to) 3+3 (to 24). Filaments free or slightly connate at base, free from tepals. Anthers basifixed, versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Inner staminal whorl rarely staminodial.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colpate, shed as monads, tricellular at dispersal. Exine semitectate, with columellate infratectum, finely reticulate.

Gynoecium Pistil composed of (one to) three (or four) connate paracarpous carpels; median/odd carpel adaxial (abaxial?). Ovary superior, unilocular. Style single, elongate, narrow, usually branched. Stigmas (one to) three (or four), papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal or basal-parietal (rarely basal). Ovules (one or) two to six (to numerous) per ovary, anatropous, ascending, bitegmic, tenuinucellar (pseudocrassinucellar). Micropyle endostomal. Outer integument two or three cell layers thick. Inner integument two or three cell layers thick. Parietal tissue absent. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustorium bicellular or quadricellular, coenocytic, micropylar. Embryogenesis solanad. Polyembryony sometimes present.

Fruit A loculicidal apicidal capsule with persistent calyx.

Seeds Aril usually present. Seed coat exotestal? Exotestal cells large; cell wall papillae with terminal nail-like thickenings. Endotestal cells thin-walled. Exotegmen? Endotegmen with thick cuticle, tanniniferous. Perisperm not developed? Endosperm copious, starchy. Embryo large, straight, chlorophyll? Suspensor absent. Cotyledons two. Germination phanerocotylar.

Cytology n = 5, 10, 15

DNA Intron present in plastid gene rpl2.

Phytochemistry Flavonol bisulphates (kaempferol, quercetin), cyanidin, tannins, proanthocyanidins (prodelphinidins), and pinitol present. Cyanogenic compounds, alkaloids, saponins and myricetin not found. Sulphated compounds frequently present. Ellagic acid?

Use Ornamental plants, fish poison.

Nakai in J. Jap. Bot. 18: 102. 10 Mar 1942

Genera/species 1/7

Distribution Tropical and subtropical regions in the Old World.

Fossils Unknown.

Habit Usually bisexual (rarely unisexual), usually annual (sometimes perennial) herbs.

Vegetative anatomy Phellogen absent? C₄ photosynthesis present. Kranz anatomy atriplicoid. Secondary lateral growth absent. Vessel elements? Imperforate tracheary xylem elements? Wood rays absent. Axial parenchyma? Sieve tube plastids P3cf type? Nodes unilacunar? with ? leaf traces. Calciumoxalate raphides present.

Trichomes Hairs?

Leaves Usually opposite (rarely almost verticillate), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata? Cuticular waxes? Leaf margin entire. Epidermis with calciumoxalate raphides.

Inflorescence Axillary or terminal, leaf-opposed panicle (sometimes umbel-like cyme) with dichasial partial inflorescences.

Flowers Actinomorphic, small. Hypogyny. Tepals five, with imbricate-quincuncial aestivation, sepaloid, free. Nectariferous disc flattened.

Androecium Stamens usually five or ten to 15 (rarely eight, sometimes 20 or more, in fascicles of three), alternitepalous. Filaments widened at base, free. Anthers basifixed, non-versatile, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate to polypantoporate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate or perforate, scabrate, spinulate or smooth.

Gynoecium Pistil composed of (three to) five (to ten) carpels, whorled, antetepalous, seemingly free (pseudapocarpy). Partial ovaries superior, unilocular. Stylodia short, ventral. Stigmas? Male flowers with pistillodium.

Ovules Placentation basal. Ovule one per carpel, anatropous (campylotropous?), bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument three cell layers thick. Inner integument ? cell layers thick. Parietal tissue approx. two cell layers thick. Nucellar cap approx. three cell layers thick. Megasporangial cells radially expanded. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit An achene with membranous epicarp covered by warts or prickles (sometimes with wing-like lateral processes).

Seeds Aril absent. Exotestal cells tangentially elongate. Endotesta? Exotegmic cells thickened. Endotegmen? Perisperm copious and nutritious. Endosperm poorly developed or absent. Embryo peripheral, curved around perisperm, well differentiated?, chlorophyll? Suspensor apically curved. Cotyledons two. Germination phanerocotylar?

Cytology x = 9

DNA

Phytochemistry Very insufficiently known. Betacyanins and betaxanthins present. Anthocyanins? Triterpene saponins?

Use Unknown.

Soriano in Bol. Soc. Argent. Bot. 23: 161. 29 Aug 1984

Genera/species 1/1

Distribution Argentina.

Fossils Unknown.

Habit Monoecious, annual herb. Leaf succulent.

Vegetative anatomy Phellogen? Cortex succulent. Secondary lateral growth anomalous (from concentric/successive cambia). Vessel elements? Imperforate tracheary xylem element libriform fibres with bordered pits? Extraxylary fibres present. Wood rays absent from secondary xylem. Axial parenchyma? Sieve tube plastids P3cf type, with a globular central protein crystal surrounded by a subperipheral dense ring of protein filaments. Nodes 1:1? Phloem parenchyma cells with phytoferritin? Calciumoxalate druses present.

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, usually entire (sometimes absent), succulent, subterete, flattened on adaxial side, convex on abaxial side, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Leaf single-veined. Stomata paracytic or anomocytic? Cuticular waxes? Leaf margin entire. Young buds surrounded by two opposite scale-like leaves (prophylls?).

Inflorescence Male flowers numerous in a terminal, dense, spike-like, bracteate inflorescence. Female flowers four or five together in axillary fascicles (or female flowers in reality solitary axillary?) in uppermost four or five leaf axils. Male flowers with two bracts or floral prophylls (bracteoles; transverse floral prophylls absent). Each female flower sunken into cortex and enclosed by one (or two?) small bracts and two very unequally sized median? floral prophylls.

Flowers Actinomorphic, small. Hypogyny? Tepals four in male flowers, with valvate-decussate aestivation, somewhat petaloid, membranous, whorled, free or connate at base. Female flowers without tepals.

Androecium Stamens four, alternitepalous. Filaments filiform, free from each other and from tepals. Anthers dorsifixed to subbasifixed, protruding, versatile, tetrasporangiate, extrorse, poricidal (dehiscing by pores due to contraction of connective). Endothecial anticlinal cell walls with frame-like thickening. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains hexaporate, cuboid (with one pore on each side), shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum (fusions?), punctate, microspinulate.

Gynoecium Pistil composed of three connate carpels; median adaxial carpel fertile. Ovary superior, unilocular. Style single, simple. Stigmas three, finally protruding, papillate, type? Pistillodium absent.

Ovules Placentation basal. Ovule one per ovary, anatropous (campylotropous?), bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Thin-walled and nutlike. Hardening inflorescence axis enclosing several nutlets aggregated into a dry syncarp.

Seeds Aril absent. Testa? Tegmen? Perisperm well developed and nutritious (with polygonal and unusually large starch grains when compared to other Caryophyllales), surrounded by embryo. Endosperm almost absent. Embryo annular, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = 12

DNA

Phytochemistry Very insufficiently known. Betanin (a tyrosine-derived glucose glycoside of betanidin) present. Proanthocyanidins not found.

Use Unknown.

Christenhusz in Phytotaxa 181: 240. 8 Oct 2014

Genera/species 1/8

Distribution Eastern and southern Africa, Madagascar, St. Helena.

Fossils Unknown.

Habit Bisexual herbs or suffrutices.

Vegetative anatomy Phellogen? Secondary lateral growth? Vessel elements? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma? Sieve tube plastids P3cf type, with central globular protein crystal surrounded by ring of protein filaments. Nodes? Calciumoxalate crystals as clusters and crystal sand.

Trichomes Hairs absent.

Leaves Alternate (spiral, often pseudoverticillate), simple, entire, linear, terete, with ? ptyxis. Stipules membranous, adnate to dilated leaf base forming a sheath; leaf sheath absent. Petiole vascular bundles? Leaf one-veined. Stomata? Cuticular wax crystalloids as rodlets. Leaf margin entire.

Inflorescence Terminal (often seemingly axillary), umbel-like thyrse.

Flowers Actinomorphic. Hypogyny. Tepals five, with imbricate aestivation, often petaloid, persistent, free. Nectaries present on adaxial side of inner staminal filaments. Disc absent.

Androecium Stamens (three to) five to 15 (to c. 30), diplostemonous. Filaments connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

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

Gynoecium Pistil composed of three to five connate antesepalous carpels. Ovary superior, trilocular to quinquelocular. Style absent. Stigmas three to five, papillate?, type? Pistillodium absent.

Ovules Placentation axile. Ovules numerous per carpel, ?-tropous, bitegmic?, crassinucellar? Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator? Parietal tissue? Megagametophyte monosporous, Polygonum type? Endosperm development ab initio nuclear, finally cellular. Endosperm haustoria? Embryogenesis solanad.

Fruit A loculicidal membranous capsule or a schizocarp with nutlike mericarps.

Seeds Aril absent. Operculum present. Seed coat? Exotesta? Endotesta? Exotegmen? Endotegmic cell walls with rod-shaped thickenings? Perisperm copious, hard, starchy. Endosperm sparse or absent? Embryo peripheral?, curved around perisperm?, well differentiated?, without chlorophyll? Cotyledons two. Radicula dorsal? Germination?

Cytology n = 8

DNA

Phytochemistry Virtually unknown. Anthocyanins present. Betalains absent?

Use Unknown.

Shipunov ex Reveal in Bot. Rev. (Lancaster) 71: 128. 20 Mai 2005

Genera/species 1/c 25

Distribution Sub-Saharan Africa, the Arabian Peninsula, southern Asia east to India.

Fossils Unknown.

Habit Bisexual, perennial herbs or suffrutices with lignified basal parts and often green photosynthesizing branches.

Vegetative anatomy Phellogen? Secondary lateral growth usually anomalous (from meristematic rings of concentric cambia in phloem or pericycle; rarely normal). Vessel elements with simple? perforation plates; lateral pits alternate? Imperforate tracheary xylem elements? Wood rays absent? Axial parenchyma? Sieve tube plastids P3c’’f type, with a central cuboidal protein crystal surrounded by a ring of protein filaments. Nodes? Calciumoxalate usually as druses.

Trichomes Hairs usually absent; glandular hairs present in some species.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation? Stomata? Cuticular waxes? Leaf margin entire.

Inflorescence Terminal, raceme-like to capitate, cymose.

Flowers Actinomorphic. Hypogyny. Sepals five, free. Petals usually five (absent in some species), free. Nectary absent? Disc absent.

Androecium Stamens (five to) seven (to ten). Filaments connate at base, adnate to tepals (epitepalous). Anthers dorsifixed, versatile?, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3(–4)-colpate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, punctate, smooth or spinulate.

Gynoecium Pistil composed of two to seven connate antesepalous carpels. Ovary superior, pseudomonomerous, two carpels being initiated: adaxial carpel sterile and much smaller than abaxial carpel, in which two ovules are developed and separated by secondary septum. Stylodia two or three or absent. Stigmas two or three, filiform, type? Pistillodium absent.

Ovules Placentation axile. Ovules one to three per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue? Megagametophyte monosporous, Polygonum type. Antipodal cells often persistent. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad.

Fruit A membranous capsule or a dimerous schizocarp with nutlike (sometimes winged) mericarps.

Seeds Dry aril/elaiosome (funicular) present or absent. Seed coat? Testa with cells in rows along dorsal suture. Tegmen? Perisperm copious, starchy. Endosperm almost absent. Embryo peripheral, curved around perisperm?, chlorophyll? Cotyledons two. Germination?

Cytology n = 9 (Limeum indicum)

DNA

Phytochemistry Virtually unknown. Anthocyanin present.

Use Unknown.

Doweld et Reveal, New Syllabus Pl. Fam.: 795. Jan 2007

Corbichoniaceae Thulin in Taxon 65(4): 790. Aug 2016

Genera/species 2/6

Distribution Sub-Saharan Africa, southern Asia.

Fossils Unknown.

Habit Bisexual, suffrutices or perennial herbs (in Corbichonia often with procumbent stems).

Vegetative anatomy Phellogen? Secondary lateral growth usually normal (sometimes anomalous from concentric meristematic rings in phloem or pericycle). Vessel elements with simple? perforation plates; lateral pits alternate? Imperforate tracheary xylem elements? Wood rays absent? Axial parenchyma? Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalates as druses or sphaeroites.

Trichomes Hairs?

Leaves Alternate (spiral), simple, entire, somewhat fleshy, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate? Stomata? Cuticular wax crystalloids as platelets. Leaf margin entire.

Inflorescence Terminal or seemingly leaf-opposite, cymose (Corbichonia), or raceme or spike arranged in groups of usually three (sometimes two) partial inflorescences (Lophiocarpus).

Flowers Actinomorphic. Hypogyny. Sepals (four or) five, with usually quincuncial aestivation, back and front sepals covering remaining ones (Corbichonia), free. Petals 15 to numerous, with valvate aestivation, free (Corbichonia), or absent (Lophiocarpus). Nectariferous disc present.

Androecium Stamens four (Lophiocarpus; occasionally seemingly three) or ten to c. 30 (Corbichonia). Filaments free from each other and from tepals. Anthers basifixed, elongate (Lophiocarpus), non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory (bicellular or tricellular). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, perforate to finely punctate, spinulate.

Gynoecium Pistil composed of two (Lophiocarpus) or five (Corbichonia) connate antesepalous carpels. Ovary superior, unilocular (Lophiocarpus) or quinquelocular (Corbichonia). Style single, simple, very short (Corbichonia), or absent (Lophiocarpus). Stigmas four (Lophiocarpus), type? Pistillodium absent.

Ovules Placentation axile (Corbichonia) or basal (Lophiocarpus). Ovule one per ovary (Lophiocarpus) or numerous per carpel (Corbichonia), campylotropous (Lophiocarpus), bitegmic, thinly crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue two or three cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad (Corbichonia) or caryophyllad (Lophiocarpus).

Fruit An achene or one-seeded drupe (Lophiocarpus) or a loculicidal capsule (Corbichonia).

Seeds Funicular aril present (Corbichonia) or absent. Seeds with small strophiole (Corbichonia). Seed coat? Exotestal cells radially elongate. Endotesta? Tegmen? Perisperm copious and nutritious. Endosperm almost absent. Embryo peripheral, curved around perisperm, chlorophyll? Cotyledons two. Germination?

Cytology n = ?

DNA

Phytochemistry Very insufficiently known. Anthocyanins and saponins present in Corbichonia. Betalains absent.

Use Unknown.

Christenhusz in Phytotaxa 181: 240. 8 Oct 2014

Genera/species 1/9

Distribution Australia.

Fossils Unknown.

Habit Bisexual, shrubs or suffrutices with lignified basal parts and often green photosynthesizing branches.

Vegetative anatomyPhellogen? Secondary lateral growth anomalous (from meristematic cylinders of concentric cambia in phloem or pericycle). Vessel elements with simple? perforation plates; lateral pits alternate? Imperforate tracheary xylem elements? Wood rays absent? Axial parenchyma? Sieve tube plastids P3c’’fs type. Nodes 1:1? Calciumoxalate usually as druses.

Trichomes Hairs usually absent.

Leaves Alternate (spiral), simple, entire, often reduced and scale-like, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation? Stomata? Cuticular waxes? Leaf margin entire.

Inflorescence Terminal, raceme-like to capitate, cymose.

Flowers Actinomorphic, often small. Hypogyny. Sepals five, free. Petals five, free, or absent. Nectary absent? Disc absent.

Androecium Stamens 4+4. Filaments connate at base, adnate to tepals (epitepalous). Anthers dorsifixed, versatile?, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3(–4)-colpate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, punctate, smooth or spinulate.

Gynoecium Pistil composed of three to seven connate antesepalous carpels. Ovary superior, with two carpels initiated: adaxial carpel sterile and much smaller than abaxial carpel, in which two ovules develop and are separated by secondary septum. Stylodia two or three or absent. Stigmas two or three, filiform, type? Pistillodium absent.

Ovules Placentation basal-axile. Ovules one to three per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue? Megagametophyte monosporous, Polygonum type. Antipodal cells usually ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A membranous loculicidal capsule.

Seeds Dry aril/elaiosome (funicular) present. Testa with cells in rows along dorsal suture. Tegmen? Perisperm copious, starchy. Endosperm almost absent. Embryo peripheral, curved around perisperm?, chlorophyll? Cotyledons two. Germination?

Cytologyn = ?

DNA

Phytochemistry Very insufficiently known. O-glycosylflavonoids and anthocyanins present. Betalains not found.

Use Unknown.

Schäferhoff et Borsch in Willdenowia 39: 223. Dec 2009

Genera/species 1/9

Distribution Mexico, Central America, the West Indies.

Fossils Unknown.

Habit Bisexual, usually annual herbs (occasionally perennial and somewhat lignified at base).

Vegetative anatomy Phellogen? Secondary lateral growth usually normal (sometimes anomalous from concentric meristematic cylinders in phloem or pericycle). Vessel elements with simple? perforation plates; lateral pits alternate? Imperforate tracheary xylem elements? Wood rays absent? Axial parenchyma? Sieve tube plastids P3f type without central protein crystal and with central starch grain. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalates as druses or sphaeroites or absent?

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, entire, somewhat fleshy, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate? Stomata? Cuticular wax crystalloids as platelets. Leaf margin entire.

Inflorescence Terminal or seemingly leaf-opposite, thyrsoid, with racemes or spikes arranged in groups of usually three (sometimes two) partial inflorescences. Floral prophylls (bracteoles two or absent).

Flowers Actinomorphic. Hypogyny. Tepals (four or) five, with quincuncial aestivation, free. Nectariferous disc present.

Androecium Stamens (two to) five to eight (or nine), alternitepalous. Filaments free from each other and from tepals. Anthers basifixed to dorsifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by apical slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 9–29-polypantoporate, shed as monads, tricellular? at dispersal. Exine tectate, with columellate infratectum, perforate to finely punctate, microspinulate.

Gynoecium Pistil composed of two to five connate carpels (with various orientation). Ovary superior, unilocular (without traces of a suture). Stylodia two to five. Stigmas two to five, papillate, type? Pistillodium absent.

Ovules Placentation free central. Ovule one per ovary, basal, campylotropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A muricate to spiny achene.

Seeds Funicular aril present. Testa crustaceous. Tegmen? Perisperm copious and nutritious. Endosperm almost absent. Embryo peripheral, curved around perisperm, chlorophyll? Cotyledons two. Germination?

Cytology n = ?

DNA

Phytochemistry Virtually unknown. Anthocyanins? Betalains?

Use Medicinal plants.

Bartling in F. G. Bartling et H. L. Wendland, Beitr. Bot. 2: 158. Dec 1825 [’Mollugineae’], nom. cons.

Pharnaceaceae Martinov, Tekhno-Bot. Slovar: 477. 3 Aug 1820 [’Pharnaceae’]; Glinaceae Mart., Consp. Regn. Veg.: 64. Sep-Oct 1835 [’Glinoideae’]; Adenogrammaceae (Fenzl) Nakai in J. Jap. Bot. 18: 101. 10 Mar 1942; Polpodaceae (Fenzl) Nakai in J. Jap. Bot. 18: 109. 10 Mar 1942

Genera/species 11/c 90

Distribution Mainly tropical and subtropical regions, with their highest diversity i southern Africa, some species in warm-temperate areas.

Fossils Unknown.

Habit Usually bisexual (in Mollugo ulei dioecious), usually annual or perennial herbs (sometimes suffrutices). Often somewhat succulent.

Vegetative anatomy C₄ photosynthesis (‘Hypertelis’) or intermediates between C₃ and C₄ photosynthesis sometimes present. Phellogen? Secondary lateral growth usually anomalous (from concentric cambia as successive meristematic cylinders in phloem or pericycle) or absent. Pericyclic fibres present. Vessel elements with simple perforation plates; lateral pits alternate?, bordered pits. Imperforate tracheary xylem elements? Wood rays usually absent. Axial parenchyma? Sieve tube plastids P3cf type, with a central globoid protein crystal surrounded by a ring of protein filaments (in Glinus P3cfs type, with starch grains). Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate usually as single crystals, lumps or druses (rarely as crystal sand, rhomboidal crystals or raphides).

Trichomes Hairs usually absent (in Glinus stellate hairs; in Mollugo ulei glandular hairs).

Leaves Usually alternate (spiral), often as a basal rosette or pseudoverticillate (sometimes opposite), simple, entire, often somewhat succulent or ericoid, with ? ptyxis. Stipules large, membranous and sometimes lobed, small or absent; leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually anomocytic (sometimes paracytic, diacytic or anisocytic?). Cuticular wax crystalloids in as platelets or rodlets. Leaf margin entire.

Inflorescence Usually terminal or seemingly axillary, cymose of various shape (in Polpoda axillary, few-flowered, cymose, or flowers solitary). Prophylls often prominent.

Flowers Actinomorphic, small. Usually hypogyny (in Coelanthum half epigyny). Tepals usually five (in Polpoda four), with imbricate aestivation, usually sepaloid (sometimes petaloid), persistent, usually free (in Coelanthum connate at base). Petaloid staminodia five to c. 20 or absent. Nectaries usually on adaxial side of inner staminal filaments (absent in Polpoda). Nectariferous disc annular or absent.

Androecium Stamens (two to) four to ten (to c. 30, in fascicles), haplostemonous or diplostemonous, usually antetepalous (in Polpoda alternitepalous) or – when more than five – in alternitepalous and epitepalous whorls. Filaments widened at base and usually connate into a tube, free from tepals. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia petaloid or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (sometimes tetracolpate; in Mollugo sometimes polypantocolpate or polypantoporate), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, tubuliferopunctate, scabrate-punctate or spinulate.

Gynoecium Pistil composed of usually two (in, e.g., Polpoda) to five (rarely more than five) connate carpels (in Adenogramma a single carpel); carpels antetepalous or median carpel adaxial. Ovary usually superior (rarely semi-inferior), usually bilocular to quinquelocular (in Adenogramma unilocular, monomerous). Style single, simple, or stylodia two to five, short, usually free (in Polpoda connate at base), or absent. Stigma single, bilobate to quinquelobate, or stigmas two to five, adaxially papillate, type? Pistillodium absent?

Ovules Placentation axile. Ovule one seemingly basal (in Adenogramma, Polpoda and Psammotropha) to numerous per carpel or ovary, (hemi)anatropous to (ana)campylotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Obturator funicular. Parietal tissue? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear, finally cellular. Endosperm haustoria? Embryogenesis solanad.

Fruit A loculicidal capsule or dehiscing by transverse slits (in Adenogramma a nutlet).

Seeds Aril present or absent. Funicle short. Operculum sometimes present. Seed coat exotestal-endotegmic. Exotestal cells indistinguished in shape. Endotesta and exotegmen crushed. Endotegmic cell walls with rod-shaped thickenings. Perisperm copious, hard, starchy. Endosperm sparse or absent. Embryo peripheral, curved around perisperm, well differentiated, without chlorophyll. Cotyledons two. Radicula dorsal. Germination?

Cytology n = 9, 18 (Glinus) – Polyploidy occurring in Mollugo s.lat. (n = 18, 27, 32, 36).

DNA

Phytochemistry Insufficiently known. Flavonols (kaempferol?, quercetin?), C-glycosyl-flavonoids, proanthocyanidins, anthocyanins, hopane saponins, and cyanogenic compounds present. Betalains not found in investigated species.

Use Medicinal plants, vegetables.

Phylogeny (simplified) of Molluginaceae based on DNA sequence data (Christin & al. 2010).

Phylogeny (simplified) of Molluginaceae based on DNA sequence data (Thulin & al. 2016).

Rafinesque in Ann. Gén. Sci. Phys. Bruxelles 5: 349. Jul-Sep 1820 [’Montidia’]

Hectorellaceae Philipson et Skipworth in Trans. Roy. Soc. New Zealand Bot. 1: 31. 20 Sep 1961

Genera/species c 10/285–290

Distribution Cosmopolitan except polar areas (mainly temperate regions on the Northern Hemisphere), Australia, Tasmania, New Zealand, Kerguélen, North and South America, with their largest diversity in western North America, western South America and southern Australia.

Fossils Unknown.

Habit Usually bisexual (rarely unisexual), perennial or annual herbs (rarely suffrutices), often succulent. Roots sometimes tuberous. Certain species? of Lewisia (’Erocallis’) with corm. Stem frequently absent. Some species are xerophytic.

Vegetative anatomy Mycorrhiza absent. At least some genera with CAM photosynthesis. At least some species with stomata present in stem epidermis. Phellogen ab initio superficial, present near stem apex. Precocious or delayed initiation of stem periderm; periderm usually absent. Endodermis sometimes prominent, with casparian dots (Montia). Primary vascular tissue usually a cylinder of bundles. Cortical and medullary parenchyma early degenerating. Secondary lateral growth normal, poor (without concentric cambia or inner phloem), or absent. Vessel elements with simple perforation plates; lateral pits alternate, pseudoscalariform, or intermediate between pseudoscalariform and helical thickenings. Imperforate tracheary xylem elements libriform fibres; non-septate. Wood rays multiseriate or absent. Axial parenchyma? Thick-walled pericyclic extraxylary phloem fibre caps absent. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Sclereids absent. Stem usually without mucilaginous idioblasts. Tanniniferous cells absent. Phloem parenchyma cells with phytoferritin? Crystal sand or druses of calciumoxalate often present; stem epidermis often with calciumoxalate crystals. At least some genera with CAM photosynthesis.

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

Leaves Alternate (spiral) or opposite, simple, entire, often perfoliate, often fleshy, with ? ptyxis. Stipules and leaf sheath absent. Leaves without paired axillary hairs, bristles or scales. Petiole vascular bundles? Venation pinnate? Stomata brachyparacytic (longitudinally orientated, with two subsidiary cells surrounding but not completely enclosing guard cells; sometimes parallelocytic or anomocytic). Cuticular wax crystalloids as procumbent platelets. Mesophyll usually with mucilaginous idioblasts (absent in Claytonia and Parakeelya). Sclereids absent. Crystals abundant. Leaf margin entire.

Inflorescence Terminal or axillary, dichasia and/or monochasia (sometimes head-, spike or raceme-like), often scorpioid, or flowers solitary axillary. Floral prophylls (bracteoles) usually two or 2+2 (inner pair median; rarely three, up to nine [Lewisia] or absent), with imbricate aestivation, lateral, apical or basal, at equal or different heights, usually persistent and dry in fruit, free or connate at base (transverse floral prophylls sometimes absent). Montiopsis sometimes with trilobate involucral bracts.

Flowers Usually actinomorphic (rarely zygomorphic). Hypogyny. Tepals (two or) three to five (to seven; in Lewisia up to 19), in one or two whorls, with imbricate aestivation, petaloid, persistent or caducous, usually free (rarely connate at base), or absent. Nectariferous disc present.

Androecium Stamens usually three to five (rarely six, seven or up to c. 100; in Lewisia up to 19; in Calandrinia/Monocosmia and Calyptridium one), in one whorl, usually antetepalous (in Lyallia alternitepalous). Filaments free or connate (all together or in fascicles), free from tepals or adnate at base. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with multinucleate cells. Staminodia usually absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tri- to polyzonacolpate, tri- to polyzonacolporate, tri- to polyzonaporate, polypantoporate, -rugate or -foraminate (in ’Montiastrum’ tholate, polypantocolpate; centres of mesocolpi vaulted forming spiniferous upraised tholi), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate, spinulate.

Gynoecium Pistil composed of two to four (to nine) connate carpels. Ovary superior, unilocular. Style single, simple, or stylodia two to four (to nine), free or connate. Stigmas capitate or lobate, papillate, Dry type. Pistillodium usually absent.

Ovules Placentation basal to free central. Ovules two to more than 100 per ovary, anatropous to amphitropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells three, ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad or solanad.

Fruit Usually a two- or three-valved capsule and/or circumscissile at base (Lewisia, Lewisiopsis; elastically or passively dehiscent; rarely poricidal; sometimes irregularly dehiscent; sometimes a one-seeded capsular utriculus, in Lenzia indehiscent or tardily dehiscent, in Philippiamra irregularly dehiscent or indehiscent; in Lyallia one- or two-seeded successively disintegrating indehiscent capsules). Fruit sometimes with deciduous calyptra formed by dry perianth and staminal remnants.

Seeds Aril usually absent (in Phemeranthus carnose or chartaceous). Seed coat testal. Strophiole or elaiosome present or absent. Outer exotestal cell walls thickened, with stalactite-shaped processes. Endotesta? Tegmen? Perisperm copious, starchy. Endosperm sparse or absent. Embryo peripheral, curved and surrounding perisperm, well differentiated, without chlorophyll. Cotyledons usually two (rarely three or four). Radicula dorsal. Germination phanerocotylar.

Cytology n = 6–13 or more – Claytonia virginiana shows extreme variation in chromosome number with n = 6 to c. 95; Lyallia: n = 48.

DNA 6 bp deletion in plastid gene ndhF.

Phytochemistry Flavonols (kaempferol), cyanidins, betalains (betacyanins, betaxanthins), alkaloids, and triterpene saponins present. Ellagic acid and cyanogenic compounds not found. Free oxalates often accumulated.

Use Ornamental plants, vegetables (Claytonia perfoliata).

Phylogeny (simplified) of Montiaceae based on DNA sequence data (Applequist & Wallace 2001). Baitaria, Lyallia, Parakeelya, and Philippiamra were not included in the study. Lyallia is sister to the [Lewisia+[Montia+Claytonia]] clade in analyses by Ogburn & Edwards (2009).

Dumortier, Anal. Fam. Plant.: 14, 16. 1829 [‘Nepenthideae’], nom. cons.

Nepenthales Bercht. et J. Presl, Přir. Rostlin: 267. Jan-Apr 1820 [‘Nepenthaceae’]; Nepenthineae Link, Handbuch 1: 369. 4-11 Jul 1829; Nepenthanae Takht. ex Reveal in Phytologia 79: 71. 29 Apr 1996

Genera/species 1/170

Distribution Madagascar, the Seychelles, Sri Lanka, Assam, Southeast Asia, southeastern China, Malesia to northeastern Australia, New Caledonia.

Fossils Fossil pollen grains, which may be assigned to Nepenthes, have been found in Late Cretaceous layers in South America and Africa, and in the Eocene and the Miocene of Europe.

Habit Dioecious, usually scrambling and climbing (sometimes epiphytic) perennial herbs (rarely lignified). Carnivorous (largely utilizing excrements and other detrimental waste products from animals living in the water solution of the pitcher and feeding on prey falling into the pitcher). Often hygrophytes.

Vegetative anatomy Mycorrhiza absent. Phellogen ab initio pericyclic. Primary vascular tissue a cylinder of bundles. Medullary vascular bundles may be present; young stems with cortical bundles. Medulla, pericycle and bark tissues with large idioblasts having spiral thickenings and cap-like apices. Pith and cortex with idioblastic helical-banded fibre-sclereids. A certain amount of lateral growth may occur. Vessel elements dimorphic, with simple (sometimes vestigial scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, usually non-septate (occasionally septate). Wood rays usually uniseriate or biseriate (rarely multiseriate), homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal banded. Sieve tube plastids S type. Nodes 5–9:5–9, multilacunar with five to nine leaf traces. Older parts of secondary xylem with gum-like deposits. Silica bodies present. Crystals?

Trichomes Hairs usually multicellular (rarely unicellular), filiform, fasciculate, rosulate, dendroid, or absent. Nectariferous glands and peltate hydathodes abundant on stems, leaves and other organs. Vascularized glands absent.

Leaves Alternate (spiral), simple, entire, with abaxially circinate, involute ptyxis. Stipules absent; leaf base sheathing, wide, gradually expanding into an even broader photosynthesizing discoid part, phyllodium, distally narrowing into a partially spirally twisted climbing organ, cirrhus, probably corresponding to petiole. This part grading into a pendant pitcher- or urn-shaped, often many-coloured organ, ascidium, catching insects. Ascidium probably corresponding to lamina and provided with lid, operculum, above mouth. Below lid, outside pitcher a spur, which may correspond to leaf apex. Pitcher differentiated into a recurved border zone, peristome, with numerous nectar-secreting glands (extrafloral nectaries) and, below these on inner side, a wax-secreting slippery zone, and at base a digesting glandular zone with proteolytic enzymes and absorbing cells partially or entirely covered by epidermis. Petiole vascular bundle transection arcuate. Chlorophyllous cells at pitcher margin oxygenating solution. On ventral side of pitcher two elongate narrowly dentate wings, corresponding to leaf margins. Venation parallelodromous to palmate. Stomata anomocytic. Cuticular waxes absent? Adaxial hypodermis present. Leaf margin entire. Water-absorbing peltate glandular hairs, hydathodes, sunken into depressions on foliar surface.

Inflorescence Terminal, raceme-like or paniculate thyrse or botryoid. Bracts and floralp prophylls (bracteoles) absent.

Flowers Actinomorphic, small. Hypogyny. Tepals (sepals?) (three or) four, with imbricate decussate aestivation, with large flat adaxial nectariferous glands, usually free (sometimes connate at base). Petals? absent. Tepal nectaries present. Disc absent.

Androecium Stamens (four to) eight to 24. Filaments connate into a central column, free from tepals. Anthers basifixed to subbasifixed?, connivent into a tube, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains inaperturate or with indistinct apertures, shed as tetrads, tricellular at dispersal. Exine tectate, with columellate infratectum, spinulate or psilate.

Gynoecium Pistil composed of (three or) four (to six) connate antesepalous carpels. Ovary superior, usually quadrilocular (rarely trilocular, quinquelocular or sexalocular). Style single, simple, very short or absent. Stigma single, capitate to discoid, papillate, Dry type. Pistillodium absent.

Ovules Placentation axile (to laminar). Ovules c. 10 to more than 50 per carpel, anatropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue one cell layer thick. Megagametophyte monosporous, Polygonum type. Chalazal projection present. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A coriaceous loculicidal capsule.

Seeds Seeds very small, numerous, fusiform, usually winged. Aril absent. Seed coat exotestal. Exotesta with strongly thickened inner cell walls. Chalaza with hairpin-shaped vascular bundle. Outer integument strongly elongating after fertilization and forming exostome. Parietal cell of ovule not further dividing. Tegmen? Perisperm not developed. Endosperm abundant, starchy, oily and proteinaceous. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two; apices with haustorial function? Germination phanerocotylar.

Cytology n = 40 – Paleopolyploidy probable.

DNA Intron present in plastid gene rpl2.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, tannins, and naphthoquinones (plumbagin, droserone, hydroxyserone) present. Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants.

de Jussieu, Gen. Plant.: 90. 4 Aug 1789 [’Nyctagines’], nom. cons.

Jalapaceae Batsch, Tab. Affin. Regn. Veg.: 224. 2 Mai 1802 [’Jalapinae’], nom. illeg.; Nyctaginales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 241. Jan-Apr 1820 [’Nyctagineae’]; Allioniaceae Horan., Prim. Lin. Syst. Nat.: 68. 2 Nov 1834 [’Allioniaceae [Nyctagineae]’]; Bougainvilleaceae J. Agardh, Theoria Syst. Plant.: 364. Apr-Sep 1858 [’Bugainvilleae’]; Pisoniaceae J. Agardh, Theoria Syst. Plant.: 363. Apr-Sep 1858 [’Pisonieae’]; Mirabilidaceae W. R. B. Oliver in Trans. Roy. Soc. New Zealand 66: 294. 1936; Nyctaginineae Nakai in J. Jap. Bot. 18: 98. 10 Mar 1942

Genera/species 31/430–470

Distribution Tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres, with their largest diversity in North and South America; Phaeoptilum in southwestern Africa.

Fossils Pollen grains assigned to Nyctaginaceae have been found in Eocene layers in Argentina, and the Late Campanian Retitricolpites multibaculates from the island of Sakhalin in Russia also resemble nyctaginaceous pollen.

Habit Usually bisexual (rarely monoecious, andromonoecious, gynomonoecious or dioecious), evergreen or deciduous trees (Leucastereae), shrubs or lianas, perennial or annual herbs. Roots sometimes fleshy or tuberous. Band-shaped sticky secretions present on internodes in Anulocaulis, Cyphomeris, and some species of Boerhavia.

Vegetative anatomy Mycorrhiza usually absent (Pisonieae may form ectomycorrhiza together with a diverse array of basidiomycetes). Many species with C₄ physiology. Phellogen usually superficial (sometimes cortical). Lateral meristem in parenchyma producing secondary cortex outwards and wood rays inwards, conjunctive tissue and a series of cambia from which isolated areas of vascular tissue (not rays) develop. Secondary lateral growth usually anomalous (usually from concentric/successive cambia). Vessel elements usually with simple (sometimes reticulate) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple or reduced bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (sometimes homocellular), or absent. Axial parenchyma apotracheal diffuse, or paratracheal vasicentric or banded (usually very scarce). Wood elements often partially storied. Phloem often intraxylary (as islands or concentric cylinders, sometimes diffuse). Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by ring of protein filaments. Endodermis in Boerhaavia with thick cell walls. Nodes 1:1?, unilacunar with one? leaf trace, often swollen. Epidermis sometimes with tanniniferous idioblasts. Heartwood often with resins etc. Calciumoxalate raphides abundant (absent in Leucastereae); styloids and/or prismatic or elongate crystals present; crystal sand present or absent.

Trichomes Hairs usually multicellular, uniseriate, or absent (sometimes dendritic or stellate; in Boldoa barbed hairs; Leucastereae with stellate or lepidote hairs); usually uniseriate glandular hairs (rarely branched) present in some species.

Leaves Usually opposite (sometimes alternate, spiral, rarely verticillate), simple, entire or lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation indistinct. Stomata usually anomocytic or paracytic (rarely actinocytic). Cuticular wax crystalloids usually absent (in some species as platelets or granules). Mesophyll usually with calciumoxalate raphides. Leaf margin usually entire (rarely serrate).

Inflorescence Terminal or axillary, panicle, spike- or umbel-like etc., or flowers solitary axillary. Partial inflorescences or single flowers often in pseudanthia, surrounded by free or connate sepaloid or petaloid bracts forming an involucre.

Flowers Usually actinomorphic (in species of Colignonia and Allionia zygomorphic). Hypogyny. Tepals (three or) four or five (to seven), in a single whorl, with induplicate-valvate or contorted (plicate?) aestivation, usually petaloid, with lower part usually carnose or coriaceous, connate into a tube or infundibuliform to campanulate, with upper part usually early caducous and lower part usually persistent around fruit. Nectary usually absent (sometimes present on receptacle). Disc present (often annular and enclosing ovary) or absent.

Androecium Stamens one to ten (to c. 40), in one or sometimes two whorls, usually alternisepalous. Filaments usually connate at base (in Leucastereae free), usually free from tepals. Anthers basifixed to dorsifixed, often versatile, tetrasporangiate, latrorse or partially introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Female flowers in Pisonieae with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–4)-colpate (Leucastereae), 6–18-pantocolpate (Belemia, Phaeoptilum) or polypantoporate (some species with twelve to numerous pores), shed as monads, tricellular at dispersal. Some species (e.g. in Mirabilis) with pollen grains having a diameter of at least 200 μm (some of the largest among angiosperms). Exine tectate or semitectate, with columellate infratectum, reticulate or punctate to anulopunctate, spinulate, echinate or tegillate.

Gynoecium Pistil composed of a single carpel. Ovary superior, unilocular (monomerous), sometimes stipitate, often with subepidermal cell layer containing numerous calciumoxalate raphides. Style usually single, simple, apical or lateral (rarely absent). Stigma linear, capitate, lobate, fimbriate, penicillate etc., papillate, Dry type. Pistillodium?

Ovules Placentation basal. Ovule one per ovary, hemianatropous or (following fertilization) anacampylotropous, ascending, usually bitegmic (in Abronia and Boerhavia unitegmic), crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte usually monosporous, Polygonum type (in species of Mirabilis Tridax type: nuclei arranged according to the principle 1:2:1). Antipodal cells usually ephemeral (often persistent, long-lived and somewhat proliferating). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad. Polyembryony present in Boerhavia.

Fruit A thin-coated nutlet or achene (often drupaceous), usually surrounded by accrescent basal part of calyx forming carnose, coriaceous or lignified diclesium ’anthocarp’ (sometimes covered with viscid secretion from glandular hairs; sometimes with prickles, warts, ridges, wings, or other supraepidermal processes, in Allionia boat-shaped with two rows of inwardly directed teeth; perianth in Leucastereae and Boldoeae persistent but not accrescent and ’anthocarp’ usually absent; fruits rarely free). Pericarp weakly developed. ’Okenia’ (nested in Boerhavia) with geocarpy.

Seeds Aril absent. Seed coat testal. Testa in Pisonia multiplicative (on raphe side), unstructured. Exotesta up to seven cell layers thick. Endotesta rarely (in Mirabilis) thickened. Tegmen? Perisperm usually abundant, usually mealy (rarely gelatinous). Endosperm sparse, dome-shaped above radicula, or absent. Embryo straight or curved around perisperm, well differentiated, with chlorophyll. Cotyledons usually two (in Abronia one), sometimes (e.g. Pisonia) unequally sized. Germination phanerocotylar.

Cytology n = 8, 10, 11, 13–17, 20, 21, 26, 27, 29, 33, 40, 42, 44, 46, 47, 56, 58, 68 – Polyploidy and aneuploidy frequently occurring. Nuclei with protein bodies?

DNA Intron absent (lost) from plastid gene rpl2. 210 bp deletion in plastid genome.

Phytochemistry Flavonols (kaempferol, quercetin), flavone-C-glycosides, cyanidin, alkaloids, triterpene saponins, and betalains (betacyanins, betaxanthins) present. Ellagic acid and cyanogenic compounds not found. Free oxalates often accumulated.

Use Ornamental plants, vegetables (Pisonia), medicinal plants, bird lime (Pisonia).

Cladogram of Nyctaginaceae based on DNA sequence data (Douglas & Manos 2007; Douglas & Spellenberg 2010).

Agardh, Aphor. Bot.: 221. 13 Jun 1824 [’Petivereae’]

Rivinaceae C. Agardh, Aphor. Bot.: 218. 13 Jun 1824 [’Rivineae’]; Petiveriales Link, Handbuch 1: 392. 4-11 Jul 1829 [‘Petiveriaceae’]; Riviniales C. Agardh in C. F. P. von Martius, Consp. Regn. Veg.: 16. Sep-Oct 1835 [‘Riviaceae’]; Hilleriaceae Nakai in J. Jap. Bot. 18: 99. 10 Mar 1942; Rivinineae Nakai in J. Jap. Bot. 18: 99. 10 Mar 1942; Seguieriaceae Nakai in J. Jap. Bot. 18: 99. 10 Mar 1942

Genera/species 9/24

Distribution Southern United States to tropical South America, the West Indies, easternmost Australia, Melanesia.

Fossils Unknown.

Habit Usually bisexual (in Ledenbergia and Monococcus dioecious), evergreen trees (in Seguieria with spines), shrubs or lianas, perennial herbs (sometimes with lignified base).

Vegetative anatomy Phellogen ab initio subepidermal. Medulla with or without diaphragms. Secondary lateral growth at least sometimes anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate, simple and/or bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple pits, usually non-septate (in Rivina?) (also vasicentric fibres). Wood rays usually multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal vasicentric scanty or banded. Intraxylary (concentric) phloem present. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate as prismatic crystals, styloids and elongate crystals (in Gallesia and Seguieria hexagonal crystals).

Trichomes Hairs usually unicellular or multicellular, uniseriate.

Leaves Alternate (spiral), simple, entire, with conduplicate ptyxis. Stipules present (rarely as tubercles or prickles) or absent; leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids as platelets. Abaxial domatia as hair tufts (in Gallesia). Epidermis often with mucilaginous idioblasts. Leaf margin entire. Extrafloral nectaries sometimes (Petiveria) present in leaf axils.

Inflorescence Terminal or axillary, raceme och spike, or cymose (in Seguieria panicle or raceme-like; in Gallesia spicate raceme or compound spike). Floral prophylls (bracteoles) usually lateral (sometimes slightly adaxial or abaxial). Extrafloral nectaries sometimes (Petiveria) present.

Flowers Usually actinomorphic (in Hilleria somewhat zygomorphic), small. Usually hypogyny (rarely epigyny or half epigyny). Tepals in various numbers, usually four (in Monococcus and Petiveria four diagonal, in Seguieria five orthogonal), with imbricate aestivation, sepaloid or petaloid, whorled, usually persistent, usually free (in Hilleria three tepals somewhat connate). Nectary absent. Disc absent.

Androecium Stamens four to c. 65 (in Rivina four, in Hilleria four to 13, in Seguieria up to c. 65), in one or two whorls. Filaments free or somewhat connate at base, free from tepals. Anthers dorsifixed, sometimes versatile, tetrasporangiate, introrse or extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers in Ledenbergia with four to six staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate (Gallesia, Seguieria) to 7–17-polypantoporate (Monococcus, Petiveria, Schindleria), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate or perforate, scabrate, spinulate or smooth.

Gynoecium Carpel usually one (rarely several, secondarily free or connate). Ovary usually superior (rarely inferior or semi-inferior), usually unilocular (apocarpy, monomerous). Style single, simple, in Seguieria and Galleria flattened and wing-like, in Rivina and Monococcus short, in Trichostigma, Schindleria, Ledenbergia and Petiveria short or absent. Stigmas one to four, plumose, penicillate (Trichostigma) or capitate (Rivina), in Seguieria and Gallesia laterally decurrent, type? Pistillodium?

Ovules Placentation basal. Ovule one per carpel, campylotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue two to 18 cell layers thick. Hypostase present. Nucellar cap approx. two cell layers thick. Nucellar beak present in Petiveria. Apical cells of megasporangium often radially elongate. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A one-seeded berry (Rivina, Trichostigma), a nutlet (in Petiveria with bristle-like processes) or a samara (in Seguieria and Gallesia with enlarged persistent lignified sepals; in Schindleria, Ledenbergia and Hilleria a utriculus; in Ledenbergia with wing-like sepals). Pericarp usually adnate to seed.

Seeds Aril absent. Seed coat exotestal, sometimes thin. Endotesta and tegmen usually collapsed. Perisperm copious and nutritious (nutrient tissue absent in mature seed in, e.g., Gallesia and Seguieria). Endosperm almost absent. Embryo peripheral, curved around perisperm, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = 18 (Hilleria, Petiveria), 36 (Petiveria, Trichostigma), 54 (Rivina) – Nuclei with protein bodies?

DNA 210 bp deletion present in plastid genome.

Phytochemistry Very insufficiently known. Flavone-C-glycosides (Trichostigma), betalains (betacyanin, betaxanthin) and sterols present. Flavonols? Alkaloids? Triterpene saponins? Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Gallesia with a garlic-like smell. Free oxalates accumulated?

Use Ornamental plants, medicinal plants.

Cladogram of Petiveriaceae based on DNA sequence data (Brockington & al. 2013).

Takhtajan in Bot. Žurn. 70: 1692. 13 Dec 1985

Physenales Takht., Divers. Classif. Fl. Pl.: 168. 24 Apr 1997

Genera/species 1/2

Distribution Madagascar.

Fossils Unknown.

Habit Dioecious, evergreen shrubs or small to medium-sized trees.

Vegetative anatomy Phellogen ab initio subepidermal. Primary vascular tissue a cylinder, without separate vascular bundles. Continuous sclerenchyma cylinder surrounding vascular cylinder in young stems. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with small simple pits or reduced bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate, homocellular or heterocellular (consisting of erect or square cells). Axial parenchyma apotracheal diffuse, paratracheal aliform, confluent, or unilateral. Sieve tube plastids? Nodes 1:1, unilacunar with one leaf trace. Pericyclic sclereids present. Secondary phloem with brachysclereids. Medullary parenchyma with sclereids. Crystals usually absent (rarely a single crystal per cell); silica grains absent. Prismatic crystals sometimes present in wood ray cells.

Trichomes Hairs absent.

Leaves Alternate (distichous), simple, entire, coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole articulated? Petiole vascular bundles arcuate. Venation pinnate, brochidodromous. Stomata anomocytic. Cuticular waxes? Mesophyll cells with calciumoxalate druses. Leaf margin entire.

Inflorescence Axillary, few-flowered raceme, or flowers solitary axillary.

Flowers Actinomorphic, small. Hypogyny. Tepals five to nine, with somewhat imbricate aestivation, sepaloid, persistent, with multicellular simple hairs inside, free or somewhat connate at base. Nectary absent. Disc absent.

Androecium Stamens (eight to) ten to 14 (to 25), in one whorl. Filaments filiform, free or partially connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum? Staminodia absent.

Pollen grains Microsporogenesis? Pollen grains 3(–5)-colpate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, echinate or spinulate, traversed by numerous microchannels.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular (with incomplete septum) at base and apex, unilocular in middle. Stylodia two, long, filiform, usually connate at base. Stigmatic surface papillate, decurrent almost the entire length of stylodia, type? Male flowers with rudimentary pistillodium.

Ovules Placentation (sub)basal to axile; placental vascular bundles inverted. Ovules two per carpel, campylotropous, ascending, bitegmic, weakly crassinucellar. Micropyle endostomal. Outer integument? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum-type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A dry one-seeded, somewhat inflated, capsular nutlike (almost drupaceous) fruit with persistent calyx.

Seeds Seed large. Aril absent. Seed coat vascularized, 16 to 20 cell layers thick, with insignificantly thickened cell walls. Exotesta? Endotesta? Tegmen? Perisperm not developed. Endosperm absent. Embryo straight, chlorophyll? Cotyledons two, unequally sized. Germination?

Cytology n = ?

DNA

Phytochemistry Virtually unknown. Cytotoxic 16-β-[(D-xylopyranosyl)oxy]oxohexadecanyl triterpene glycosides and oxohexadecanoic acid present. Ellagic acid? Alkaloids?

Use Unknown.

Brown in J. H. Tuckey, Narr. Exped. Zaire: 454. 5 Mar 1818 [’Phytolaceae’], nom. cons.

Phytolaccales Link, Handbuch 1: 390. 4-11 Jul 1829 [‘Phytolacceae’]; Sarcocaceae Raf., Fl. Tellur. 3: 55. Nov-Dec 1837 [’Sarcocidia’]; Phytolaccineae Engl., Syllabus, ed. 2: 112. Mai 1898

Genera/species 3–4/30–35

Distribution Tropical and subtropical regions, with their largest diversity in South America.

Fossils Unknown.

Habit Usually bisexual (rarely dioecious), evergreen trees, shrubs or lianas (Ercilla), perennial or annual herbs (Anisomeria is a succulent). Anisomeria and some species of Phytolacca with napiform roots.

Vegetative anatomy Mycorrhiza usually absent. Phellogen ab initio subepidermal. Medulla in Phytolacca sometimes septated by diaphragms. Secondary lateral growth normal or anomalous (from concentric/successive inner-cortical cambia). Vessel elements with simple perforation plates; lateral pits alternate, non-bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple or reduced bordered pits, septate (also vasicentric fibres). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse. Intraxylary phloem present in Phytolacca dioica. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate styloids and raphides present.

Trichomes Hairs usually unicellular or multicellular, uniseriate.

Leaves Alternate (spiral), simple, entire, with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate; finer veins indistinct. Stomata anomocytic or paracytic. Cuticular wax crystalloids as platelets. Epidermis often with mucilaginous idioblasts. Leaf margin entire.

Inflorescence Terminal or axillary, or leaf-opposite, raceme or spike.

Flowers Usually actinomorphic (in some species of Anisomeria zygomorphic), small. Hypogyny. Tepals (four or) five, in, e.g., Phytolacca spiral, usually with quincuncial (in Anisomeria descending-cochlear) aestivation, sepaloid, fleshy, usually persistent, free. Nectariferous disc absent in Anisomeria.

Androecium Stamens five to c. 30. Filaments usually connate at base (sometimes free), free from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers sometimes with staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate or perforate, scabrate, spinulate or smooth.

Gynoecium Carpels three to 16, seemingly or secondarily free or connate below and with free stylodia (carpels in Nowickea on gynophore), initiated in a ring around receptacular apex, alternitepalous or antetepalous. Ovary single, superior, unilocular (pseudapocarpy). Stylodia three to 16, free or connate, more or less gynobasic. Stigmas free, type? Male flowers sometimes with pistillodium.

Ovules Placentation basal or subbasal. Ovule one per carpel, campylotropous (amphitropous?), apotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator present. Hypostase present. Parietal tissue approx. two cell layers thick. Nucellar cap massive. Apical cells of megasporangium radially elongate. Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes slightly proliferating. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad, caryophyllad or chenopodiad.

Fruit An assemblage of one-seeded berries or a berry-like syncarpous fruit.

Seeds Aril absent. Seed coat exotestal. Exotestal cells thick-walled? Endotesta and tegmen largely collapsed. Perisperm copious and nutritious. Endosperm poorly developed or absent. Embryo peripheral, curved around perisperm, well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = 9, 18, 36 (Phytolacca) – Nuclei with protein bodies?

DNA 210 bp deletion present in plastid genome.

Phytochemistry Flavonols (kaempferol), betalains (betacyanins, betaxanthins), alkaloids, triterpene saponins, sterols, and pinitol present. Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Free oxalates accumulated.

Use Ornamental plants, dyeing sources (Phytolacca), medicinal plants.

de Jussieu, Gen. Plant.: 92. 4 Aug 1789 [’Plumbagines’], nom. cons.

Staticaceae Cassel, Lehrb. Nat. Pflanzenordn.: 215. Apr-Mai 1817 [’Staticeae’]; Plumbaginales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 241. Jan-Apr 1820 [‘Plumbagineae’]; Staticales Link, Handbuch 2: 262. 4-11 Jul 1829 [‘Staticinae’]; Armeriaceae Horan., Prim. Lin. Syst. Nat.: 68. 2 Nov 1834; Plumbaginopsida Endl., Gen. Plant.: 346. Dec 1837 [’Plumbagines’]; Plumbaginineae J. Presl in Nowočeská Bibl. [Wšobecdný Rostl.] 7: 1238. 1846[‘Plumbagines‘]; Limoniaceae Ser., Fl. Pharm.: 456. 1851, nom. cons.; Aegialitidaceae Lincz. in Novosti Sist. Vyssik Rast. [Nov. Syst. Plant. Vasc.] 1968: 173. 18 Dec 1968; Plumbaginanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992; Plumbaginidae C. Y. Wu in Acta Phytotaxon. Sin. 40: 291. 2002

Genera/species 28/625–835

Distribution Cosmopolitan except Antarctica, with their largest diversity in arid and saline environments in the Mediterranean and Southwest to Central Asia.

Fossils Uncertain.

Habit Bisexual, usually perennial herbs or shrubs (sometimes annual herbs, rarely lianas). Many species are xerophytes or halophytes.

Vegetative anatomy Phellogen ab initio subepidermal, later cortical. Cortical and medullary bundles often present. Secondary lateral growth in Acantholimon, Aegialitis and Limoniastrum anomalous (often from successive/concentric cambia). Vessel elements with simple perforation plates; lateral pits alternate, simple pits? Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate? (also vasicentric tracheids). Wood rays multiseriate, homocellular or absent. Axial parenchyma paratracheal scanty. Wood elements sometimes storied. Intraxylary phloem present in some species. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Silica bodies often abundant. Crystals?

Trichomes Hairs unicellular or multicellular, uniseriate; glands stalked or unstalked (sometimes shaggy). Vascularized mucilage glands present.

Leaves Alternate (spiral), simple, entire or pinnately lobed, usually herbaceous (rarely scale-like; in Aegialitis coriaceous), with convolute, involute or flat ptyxis. Petiole vascular bundle transection arcuate. Stipules usually absent (rarely well developed); leaf sheath usually absent (present in Aegialitis?). Venation usually pinnate (sometimes palmate; in Aegialitis parallelodromous). Stomata anomocytic, anisocytic or paracytic (Ranunculaceae or Rubiaceae type). Cuticular wax crystalloids usually absent (sometimes as irregular platelets). Epidermis often with chalk glands secreting water and calcium salts; epidermal salt-excreting glands present in halophytes. Extrafloral nectaries sometimes present on abaxial side of midvein in Plumbaginoideae. Mesophyll often with secretory glands; abaxial vein axils often with upraised vascularized mucilage glands; in Staticoideae with sclerenchymatous idioblasts (with branched sclereids; sclerenchyma in Limonium with asterosclereids, osteosclereids, filiform sclereids, etc.). Leaf margin sinuate, crenate or entire.

Inflorescence Terminal, racemose (Plumbaginoideae) or cymose (Staticoideae), simple or compound; capitate inflorescence in Armeria composed of glomera/drepania consisting of cincinni; floral bracts aborting; glomeral bracts forming involucrum around capitulum; lower parts growing downwards forming sheath around uppermost part of peduncle. Bracts often sheathing, dry and membranous. Floral prophylls (bracteoles) in Plumbaginoideae often with extrafloral nectaries.

Flowers Actinomorphic. Hypogyny. Sepals five, with valvate or plicate aestivation, persistent, often membranous or petaloid, with five or ten ridges, usually connate into a tube (rarely free). Petals five, with usually contorted (sometimes imbricate) aestivation, often persistent, connate usually only at base (sometimes entirely), in Aegialitis coriaceous; petal and staminal primordia common. Nectaries often present (sometimes as five glands alternating with stamens). Disc absent.

Androecium Stamens five, alternisepalous, antepetalous. Filaments free or connate, free from petals (Plumbaginoideae) or adnate at base to petals (Staticoideae); adaxial side often with basal nectary. Anthers usually dorsifixed (in Aegialitis basifixed), sometimes versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory or amoeboid-periplasmodial. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–5)-colpate (rarely pantocolpate), tetra- or hexarugate or irregular, shed as monads, usually tricellular (sometimes bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, in Plumbaginoideae tectate and verrucate, in Staticoideae reticulate. Pollen grains in Plumbaginoideae Plumbago type (columellae very irregular, tectum continuous), in Staticoideae Armeria type (columellae straight and irregular, tectum discontinuous, reticulate). Pollen grains in Staticoideae often dimorphic.

Gynoecium Pistil composed of five connate eusyncarpous carpels. Ovary superior, unilocular. Style single, lobate at apex (Plumbaginoideae), or stylodia five, free or connate below (Staticoideae). Stigmas five, capitate to cylindrical, papillate (sometimes with multicellular papillae), Dry type. Pistillodium absent. Heterostyly frequent in Plumbaginoideae.

Ovules Placentation basal. Ovule one per ovary, anatropous or circinotropous, pendulous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Funicle long and curled. Obturator usually present between ovary apex and micropyle. Parietal tissue two or three cell layers thick. Nucellar cap in Plumbagella approx. two cell layers thick. Megagametophyte tetrasporous, 4- or 8?-nucleate, Plumbago type (Ceratostigma, Plumbago, Dyerophytum), Plumbagella type (Plumbagella), Fritillaria type (Limonium, Armeria), Adoxa type?, or Penaea type? Proendosperm nucleus triploid or tetraploid. Endosperm development ab initio nuclear. Endosperm haustorium chalazal. Embryogenesis solanad.

Fruit Usually a membranous nutlet (sometimes a capsule [denticidal?] or a pyxidium), entirely or partially enclosed by persistent calyx.

Seeds Aril absent. Seed coat exotestal-endotegmic, winged. Exotesta? Endotesta? Exotegmen? Endotegmen persistent. Perisperm not developed. Endosperm usually copious (sometimes sparse or absent), with simple starch grains and proteins, tetraploid or pentaploid. Embryo large, straight, well differentiated, with chlorophyll. Cotyledons two, flat. Germination phanerocotylar.

Cytology n = 6, 7 (Plumbaginoideae) or x = 8, 9 (Staticoideae) – Polyploidy and aneuploidy frequent (especially in Staticoideae). Agamospermy present in Limonium etc.

DNA Intron present in plastid gene rpl2. Intron in plastid gene rpl16 absent (lost) in Staticeae (e.g. Limonium gmelinii). Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavonol sulphates (cholin-O-sulphate), and gallic acid present. Hydrolyzable tannins based on ellagic acid and condensed tannins based on leucodelphinidin present in Staticoideae. Naphthoquinone (plumbagin) present in all investigated Plumbaginoideae (not found in Staticoideae). Cyanidin and prodelphinidins present in Plumbaginoideae. Alkaloids and cyanogenic compounds present in Plumbago. Ellagic acid present in Aegialitis. Glycine betaines (quaternary ammonium compounds) present in some species of Plumbago and Limonium. Saponins not found. Oxalate sometimes accumulated.

Use Ornamental plants, medicinal plants.

One of three most-parsimonious cladograms from successive weighting of Plumbaginaceae based on DNA sequence data (Lledó & al. 1998).

de Jussieu, Gen. Plant.: 82. 4 Aug 1789 [’Polygoneae’], nom. cons.

Persicariaceae Martinov, Tekhno-Bot. Slovar: 473. 3 Aug 1820 [’Persicariae’]; Polygonales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 240. Jan-Apr 1820 [‘Polygoneae’]; Rumicaceae Martinov, Tekhno-Bot. Slovar: 554. 3 Aug 1820 [’Rumoides’]; Rumicales Burnett, Outl. Bot.: 1141. Jun 1835 [‘Rumicinae’], nom. illeg.; Eriogonaceae (Dumort.) G. Don in R. Sweet, Hort. Brit., ed. 3: 580. 23 Oct 1839 [’Eriogoneae’]; Polygonopsida Brongn., Enum. Plant. Mus. Paris: xxvii, 100. 12 Aug 1843 [’Polygonoideae’]; Atraphaxidineae H. Gross in Bot. Jahrb. Syst. 49: 250. 1913; Calligonaceae Khalk. in Dokl. Akad. Nauk. Uzbeksk. SSR 1985(11): 45. Nov 1985; Polygonanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992; Polygonidae C. Y. Wu in Acta Phytotaxon. Sin. 40: 294. 2002

Genera/species 42/1.480–1.530

Distribution Cosmopolitan except Antarctica, with their largest diversity in tempererate regions on the Northern Hemisphere.

Fossils Numerous fossilized fruits have been found in Late Cretaceous (Maastrichtian) and Paleocene later layers in North America, but also from Cenozoic strata in Asia and Europe. Lower Paleocene leaf fossils, Paranymphaea crassfolia, from North America were assigned to Polygonaceae (McIver & Basinger 1993).

Habit Usually bisexual (sometimes monoecious, polygamomonoecious or dioecious), usually perennial or annual herbs (often climbing or twining) or shrubs (rarely trees or lianas). Stem and branches often sulcate, geniculate, striate and/or hollow, often with swollen nodes (branches in Muehlenbeckia and Calligonum photosynthesizing phyllocladia).

Vegetative anatomy Mycorrhiza usually absent (endomycorrhiza present in Eriogonum; ectomycorrhiza present in Coccoloba). Phellogen ab initio usually subepidermal (sometimes pericyclic). Subepidermal collenchyma or sclerenchyma strands frequently present in stem. Endodermis often significant. Lignified species with normal or anomalous secondary lateral growth (in Antigonon from successive/concentric cambia). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits sometimes present (Muehlenbeckia). Imperforate tracheary xylem elements tracheids or ? with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays usually uniseriate (sometimes multiseriate), homocellular or heterocellular (with dark-stained inclusions). Axial parenchyma apotracheal diffuse, or paratracheal scanty, vasicentric, confluent, or banded. Wood elements sometimes partially storied. Intraxylary phloem present or absent. Sieve tube plastids usually S type (sieve tube plastids with protein fibrils present in Triplarieae). Nodes 3:3, trilacunar with three leaf traces, or ≥5:≥5, multilacunar with five or more traces. Mucilage cells abundant. Calciumoxalate as prismatic crystals, crystal sand and druses often present.

Trichomes Hairs unicellular or multicellular, usually uniseriate (sometimes stellate or glandular, also peltate-lepidote).

Leaves Usually alternate (spiral; rarely opposite or verticillate), usually simple (rarely pinnately compound), entire or lobate, sometimes fleshy or coriaceous (in Muehlenbeckia and Calligonum rudimentary), usually with revolute (in Muehlenbeckia convolute) ptyxis. Stipules usually enclosing stem as a caducous or persistent, membranous, often lobate or fimbriate tubular ocrea (usually absent in the Eriogonum clade; rudimentary in some species of Chorizanthe). Colleters present in association with leaves. Foliar tendrils present in Antigonon. Petiole vascular bundle transection usually annular (sometimes D-shaped; bundles scattered in some species of Coccoloba). Extrafloral nectaries present in some species on abaxial side of petiolar pulvinus. Venation pinnate or palmate. Stomata usually anomocytic (rarely diacytic, anisocytic, helicocytic, or paracytic). Cuticular wax crystalloids as platelets or rodlets. Lamina sometimes gland-dotted. Epidermis often with mucilage cells. Leaf margin usually entire (sometimes crenate or lobed).

Inflorescence Usually terminal or axillary, simple or compound panicle, thyrsoid, or spike-, head- or raceme-like (flowers rarely single axillary). Each flower usually subtended by a persistent membranous, tubular ocreola consisting of two connate floral prophylls (bracteoles). Partial inflorescence in the Eriogonum clade surrounded by and partially enclosed by involucrum.

Flowers Actinomorphic, small. Pedicel articulated. Hypanthium present or absent. Hypogyny. Tepals usually five or 3+3 (rarely 2+2 or 3+6), with imbricate quincuncial aestivation, sepaloid or petaloid, spiral, usually persistent, connate at base. Nectariferous disc annular, inserted around ovary base, or single nectaries present between staminal bases (sometimes absent).

Androecium Stamens usually 3+3 (sometimes two, three, 2+2, five, 4+4, or 3+6), alternisepalous or antesepalous. Filaments filiform, free or connate at base, often adnate to tepal bases forming a ring. Anthers basifixed or dorsifixed, often versatile, tetrasporangiate, usually introrse (sometimes latrorse or extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, tricolporate or pantoporate, shed as monads, usually tricellular (rarely bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, punctate eller striate, often spinulate or echinulate.

Gynoecium Pistil composed of (two or) three (or four) connate eusyncarpous carpels; median carpel adaxial; when tepals 3+3, then carpels antesepalous. Ovary superior, unilocular (sometimes seemingly multilocular by secondary septa). Style single, simple, or stylodia (two or) three (or four), usually long (rarely short), separate or connate below. Stigma one or stigmas several, entire or lobate, filiform, penicillate, peltate, or capitate, papillate or non-papillate, Dry type. Pistillodium? Heterstyly occur in some genera.

Ovules Placentation basal and free central. Ovule one per ovary, usually orthotropous (occasionally campylotropous to anatropous), ascending, usually bitegmic (rarely unitegmic), crassinucellar. Funicle present (representing a reduced free central placenta?), or absent. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Nucellar beak present. Hypostase present or absent. Archespore usually unicellular (rarely multicellular). Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells usually uninucleate (in Rumex multinucleate). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit A usually triangular (rarely lenticulate) achene, sometimes winged or with bristles (rarely a berry), often surrounded by persistent and sometimes accrescent tepals (in Emex and Oxygonum surrounded by receptacle/hypanthium).

Seeds Aril absent. Seed coat testal. Exotesta? Endotesta? Tegmen? Perisperm entirely or almost absent. Endosperm copious, mealy or horny (in Coccolobeae and Triplareae ruminate), with simple starch grains, oil and proteins. Embryo usually straight to curved (rarely plicate), usually lateral to peripheral (rarely central), without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology x = (4–)7–13(–17) – Polyploidy and aneuploidy frequently occurring. Dioecious species of Rumex with a sex-determining X-autosome balance system.

DNA Intron present in the plastid gene rpl2. Mitochondrial intron coxII.i3 lost. Plastid IR expanded.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), hyperoside (quercetin 3-O-galactoside), flavonol sulphates, flavone-C-glycosides, catechins, O-methylated flavonoids, hexaoxygenated flavonoids, condensed and hydrolyzable tannins, proanthocyanidins (prodelphinidins), methylated and non-methylated ellagic acids, gallic acid, caffeic acid, indole alkaloids and other alkaloids, saponins, sesquiterpene lactones?, free soluble oxalic acid, oleanolic acid derivatives, pinitol, fagopyrine and protofagopyrine, polyacetate derived anthraquinones (oxymethyl anthraquinone etc.), and naphthoquinones and acetophenones present. Cyanogenic compounds not found.

Use Ornamental plants, vegetables (Rheum, Rumex etc.), fruits (Coccoloba uvifera), starch sources (Fagopyrum).

Cladogram of Polygonaceae based on DNA sequence data (ITS; Sanchez & al. 2009; Burke & al. 2010; Sanchez & al. 2011; Schuster & al. 2011).

Cladogram (Bayesian inference) of Eriogoneae based on DNA sequence data (Kempton 2012).

de Jussieu, Gen. Plant.: 312. 4 Aug 1789 [’Portulaceae’], nom. cons.

Portulacales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 238. Jan-Apr 1820 [‘Portulaceae’]; Portulacineae Engl., Syllabus, ed. 2: 113. Mai 1898

Genera/species 1/100–115

Distribution Cosmopolitan except polar areas, with their largest diversity in tropical and subtropical parts of North and South America.

Fossils Unknown.

Habit Bisexual, perennial or annual herbs (in Portulaca suffrutescens somewhat lignified at stem base). Roots often tuberous. Usually leaf succulents.

Vegetative anatomy Mycorrhiza usually absent. C₄ photosynthesis alternating with CAM photosynthesis; leaves with Kranz’ anatomy. Root cortex often with subelliptic thin-walled brachysclereids. Stem epidermis with or without parallelocytic stomata. Phellogen ab initio epidermal. Delayed initiation of stem periderm. Medulla sometimes with wide-band tracheids. Cortical fibres absent. Secondary lateral growth normal or absent. Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements libriform fibres with simple pits; non-septate. Wood rays multiseriate or absent. Axial parenchyma? Thick-walled pericyclic extraxylary phloem fibre caps absent. Sieve tube plastids P3cf type? Nodes 1:1, unilacunar with one leaf trace? Sclereids often present. Mucilaginous idioblasts abundant. Tanniniferous cells absent. Phloem parenchyma cells with phytoferritin? Parenchyma and epidermis often with numerous calciumoxalate crystals or druses.

Trichomes Hairs unicellular or multicellular, uniseriate; stem epidermis sometimes with papillae or multicellular hairs.

Leaves Usually alternate (rarely opposite), simple, entire, usually succulent, flat to terete, with various ptyxis? Stipules and leaf sheath absent. Leaves with or without few to numerous axillary biseriate or oligoseriate hairs or bristles (in Portulaca somalica and P. wightiana multiseriate, scale-like). Petiole vascular bundle transection arcuate? Venation pinnate. Stomata parallelocytic, usually transversely orientated. Cuticular wax crystalloids usually absent (rarely as relatively irregular platelets?). Mesophyll with mucilaginous idioblasts. Leaf margin entire.

Inflorescence Terminal, few-flowered, capitate, cymose, often surrounded by involucral bracts. Floral prophylls (bracteoles) 2+2, inner pair median. Two large floral prophylls (bracteoles) sometimes subtending flower and four smaller prophylls in a whorl separated from first pair by short internode. Transverse floral prophylls with axillary flowers, or absent; median prophylls without flowers. Sepaloid floral prophylls two.

Flowers Actinomorphic, small. Hypanthium present. Epigyny to half epigyny. Tepals (three to) five (to eight), with imbricate aestivation, petaloid, free or slightly connate at base (rarely absent). Nectary absent? Disc absent.

Androecium Stamens four to numerous. Androecial ring primordium present. Filaments free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually polypantocolpate (polyrugate), shed as monads, tricellular? at dispersal. Exine tectate, with columellate infratectum, spinulate or echinate.

Gynoecium Pistil composed of (four or) five (to eight) connate carpels. Ovary inferior or semi-inferior, ab initio multilocular, later unilocular. Stylodia (four or) five (or six). Stigmas papillate, Dry type. Pistillodium absent.

Ovules Placentation usually free central (rarely basal). Ovules numerous per ovary, anatropous?, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Apical cells of megasporangium often radially elongate. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus? Antipodal cells two?, ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad.

Fruit A pyxidium with apical operculum, dehiscing together with perianth remnants, stamens and style as a dry calyptra. Pericarp undifferentiated.

Seeds Aril hilar, spongy. Seed coat? Anticlinal testal cell walls sinuous. Tegmen? Perisperm copious, starchy. Endosperm sparse or absent. Embryo curved, well differentiated, without chlorophyll. Cotyledons two. Radicula dorsal. Germination phanerocotylar?

Cytology n = (8–)10

DNA Intron absent from plastid gene rpl2. 6 bp deletion in plastid gene ndhF. C. 500 bp deletion in plastid gene rbcL.

Phytochemistry Insufficiently known. Betalains (betacyanins, betaxanthins) and sterols present. Ellagic acid and cyanogenic compounds not found.

Use Ornamental plants, vegetables, medicinal plants.

Prance in Bull. Jard. Bot. État 38: 141. 30 Jun 1968

Rhabdodendrales Doweld, Tent. Syst. Plant. Vasc.: xli. 23 Dec 2001; Rhabdodendranae Doweld, Tent. Syst. Plant. Vasc.: xli. 23 Dec 2001

Genera/species 1/3

Distribution Tropical South America.

Fossils Unknown.

Habit Usually bisexual (rarely androdioecious), evergreen trees or shrubs.

Vegetative anatomy Phellogen? Cortical vascular bundles sometimes present. Anomalous secondary lateral growth from successive cambia present in two species (not in Rhabdodendron macrophyllum). Vessel elements with simple perforation plates; lateral pits scalariform or alternate, bordered pits. Imperforate tracheary xylem elements tracheids (libriform fibres absent) with simple or bordered pits, non-septate. Vestured pits present. Wood rays uniseriate or multiseriate, usually heterocellular (sometimes homocellular). Axial parenchyma abaxial, apotracheal sparsely diffuse, or paratracheal scanty, vasicentric, or banded. Intraxylary (concentric) phloem present. Sieve tube plastids Pcs type, with one peripheral polygonal protein crystalloid and multiple starch grains. Nodes ?:?, multilacunar with ? leaf traces. Parenchyma with resinous secretory cavities. Dark-staining substances present especially in wood rays. Sclereids present. Silica bodies sometimes present in wood. Acicular or elongate crystals, styloids, crystal sand and other types of crystals and crystal complexes (e.g. sphaerocrystals) abundant.

Trichomes Hairs multicellular, fimbriate, short-stalked, peltate.

Leaves Alternate (spiral), simple, entire, coriaceous, with revolute ptyxis. Stipules small, caducous or absent; leaf sheath absent. Petiole vascular bundles transection annular, with often separate bundles; wing bundles present; medullary bundles sometimes present. Leaf base usually widened. Venation pinnate. Stomata anomocytic, without subsidiary cells. Cuticular waxes? Lamina revolute, gland-dotted, lysigenic secretory cavities with resins, scattered lipid bodies and on abaxial side small short-stalked peltate hairs with silica inclusions. Mesophyll with sclerenchymatous idioblasts, traversed by fibre-like simple or branched sclereids representing elongations of vein endings; many mesophyll cells with silica bodies and silicified walls. Leaf margin entire.

Inflorescence Axillary or supra-axillary, simple or compound, raceme-like or racemose.

Flowers Actinomorphic. Hypanthium present. Hypogyny. Sepals five, with imbricate quincuncial aestivation, partially or entirely connate. Petals (four or) five, with cochleate, quincuncial or valvate (imbricate below) aestivation, sepaloid, gland-dotted, caducous, free. Nectary absent. Disc absent.

Androecium Stamens 27 to 53, in three whorls. Filaments flattened, very short, persistent, free from each other and from petals. Anthers basifixed, non-versatile, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits), with monocotyledonous wall development. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolp(or)ate (rarely tetracolp[or]ate), shed as monads, bicellular (or tricellular?) at dispersal. Exine tectate or semitectate, with columellate infratectum, punctate to finely reticulate.

Gynoecium Pistil composed of a single carpel. Ovary superior, unilocular. Stylodium gynobasic, thick. Stigmatic surfaces usually reaching from base or centre of abaxial side to apex (unilateral, dorsal stigma), type? Pistillodium absent?

Ovules Placentation basal. Ovules ab initio usually two, later one (one ovule degenerating) per ovary, (hemi)campylotropous, epitropous, bitegmic in micropylar end (unitegmic elsewhere), crassinucellar. Micropyle ?-stomal. Outer integument four or five cell layers thick. Inner integument two to five cell layers thick. Parietal tissue ten or more cell layers thick. Nucellar cap absent. Archespore multicellular. Megagametophyte monosporous, aberrant Polygonum type (unique variation). Endothelium not formed. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A small short-stalked dry one-seeded drupe, surrounded at base by persistent calyx/hypanthium and with swollen lignified distal part of pedicel.

Seeds Aril absent. Seed coat testal. Testa thin. Exotestal cells tangentially elongate. Endotestal cells shortly tracheidal. Tegmen? Perisperm poorly developed. Endosperm poorly developed. Embryo curved, well differentiated, with chlorophyll. Radicula curved inwards against hilum. Cotyledons two, large, fleshy. Germination cryptocotylar.

Cytology n = 10

DNA

Phytochemistry Triterpenoids, methylated and non-methylated ellagic acid, O-alkylated ellagic acid derivatives, prodelphinidin, oleanolic acid derivatives, and cyanogenic compounds present. Alkaloids not found.

Use Timber.

Behnke in Taxon 46: 503. 15 Aug 1997

Genera/species 1/1–2

Distribution Arid regions in southwestern North America.

Fossils Unknown.

Habit Monoecious (sometimes dioecious?), evergreen? shrubs with spines. Xeromorphic halophytes.

Vegetative anatomy Phellogen? Secondary lateral growth anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Non-vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple or (reduced) bordered pits, non-septate (also vasicentric tracheids). Wood rays absent. Axial parenchyma paratracheal vasicentric. Intraxylary (concentric or diffuse) phloem present. Vessel and phloem elements and/or parenchyma storied. Sieve tube plastids P3cf type, with a globular central protein crystal and a subperipheral dense ring of protein filaments. Nodes? Heartwood with gums etc. Parenchyma often with idioblasts containing elongate and rhomboidal crystals or solitary styloids.

Trichomes Hairs ’sub-bladder trichomes’ (Carolin 1983).

Leaves Alternate (spiral), simple, entire, fleshy, linear, terete, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation? Stomata? Cuticular waxes? Raphides? Leaf margin entire.

Inflorescence Male inflorescence terminal, spike- or catkin-like. Female flowers axillary, solitary or pairwise. Floral prophylls (bracteoles) absent at least in male inflorescences (in female flowers sometimes present, bilobate, pairwise connate into a tube).

Flowers Actinomorphic, small. Probably half epigyny. Tepaloid structures (bracts or sepaloid tepals?) in female flowers two, scale-like, fleshy, persistent, connivent into a tube, tepals adnate to ovary. Male flowers inserted inside stalked peltate scales, without tepals. Nectary absent. Disc absent.

Androecium Stamens usually one to three (to four), inserted below a stalked peltate scale. Filaments short, free. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains pantoporate, foraminate, with raised pore margins, shed as monads, tricellular? at dispersal. Exine tectate, with columellate infratectum, sculpturing?

Gynoecium Pistil composed of two connate carpels. Ovary inferior?, unilocular. Style single, simple, short, or absent. Stigmas two, horizontally expanded, papillate, type? Pistillodium absent.

Ovules Placentation basal. Ovule one per ovary, campylotropous?, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A nutlet surrounded by persistent and accrescent fleshy tepals, horizontally winged at apex.

Seeds Seed vertical. Aril absent. Testa membranous. Exotesta? Endotesta? Tegmen? Perisperm copious?, nutritious. Endosperm absent. Embryo peripheral, spirally twisted around perisperm, with chlorophyll. Cotyledons two. Germination?

Cytology n = 9

DNA Deletion of 210 bp in plastid genome?

Phytochemistry Insufficiently known. Betalains (betacyanins and betaxanthins) present. Oxalic acid often accumulated. Triterpene saponins? Anthocyanins not found.

Use Tools.

van Tieghem in Bot. Jahresber. (Just) 25(2): 422. 19 Jan 1900 [emend. Tiegh. ex Reveal et Hoogland, Bull. Mus. Natl. Hist. Nat. (Paris), ser. 4, B Adansonia 12: 206. 24 Nov 1990]

Simmondsiales Reveal in Novon 2: 239. 13 Oct 1992; Simmondsianae Doweld, Tent. Syst. Plant. Vasc.: xliii. 23 Dec 2001; Simmondsiineae Reveal in Kew Bull. 66: 48. Mar 2011

Genera/species 1/1

Distribution Southwestern United States, northwestern Mexico.

Fossils Unknown.

Habit Dioecious, evergreen shrubs. Xerophytes.

Vegetative anatomy Phellogen ab initio usually pericyclic. Secondary lateral growth anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate or opposite, bordered pits. Imperforate tracheary xylem elements tracheids with large bordered pits, non-septate. Wood rays rare, uniseriate or multiseriate, heterocellular. Axial parenchyma rare, apotracheal diffuse, or paratracheal scanty, or absent. Intraxylary phloem present. Sieve tube plastids Ss type, with few small starch grains. Nodes 1:1, unilocular with one leaf trace. Calciumoxalate as druses and prismatic crystals.

Trichomes Hairs short, unicellular or multicellular, uniseriate.

Leaves Opposite, simple, entire, coriaceous, articulated near leaf insertion, with flat ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate? Venation pinnate, brochidodromous. Stomata cyclocytic or laterocytic (or anomocytic?), sunken. Cuticular waxes? (organ specific). Mesophyll with calciumoxalate as druses and single prismatic crystals. Leaf margin entire.

Inflorescence Male inflorescence terminal or axillary, cymose, head-like. Female flowers usually solitary axillary (rarely fascicular in axillary racemes).

Flowers Actinomorphic, small. Hypogyny. Tepals (four or) five (or six), with imbricate aestivation, sepaloid, in male flowers fimbriate, in female flowers foliaceous, persistent. Petals absent. Nectary absent. Disc absent.

Androecium Stamens (4+4 or) 5+5 (to 8+8), diplostemonous. Filaments short, stout, free from each other and from tepals. Anthers longer than connectives, basifixed or subbasifixed, non-versatile, tetrasporangiate, extrorse (or latrorse?), longicidal (dehiscing by longitudinal slits). Tapetum secretory, in later stages with a tendency to disintegration of membranes. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate to indistinctly triporate (aperture with operculoid central part), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, punctate-scabrate, beset with small dots of tiny spinules.

Gynoecium Pistil composed of three (or four) connate carpels. Ovary superior, usually trilocular (sometimes quadrilocular). Stylodia three (or four), free, long protruding, subulate, recurved, papillate and hairy, caducous, with stylar canal. Stigmas little differentiated, papillate, type? Pistillodium absent.

Ovules Placentation apical to subapical-axile. Ovule one (or two) per carpel, anatropous, pendulous, apotropous, bitegmic, tenuinucellar (crassinucellar?). Micropyle endostomal. Outer integument six to ten cell layers thick. Inner integument three to five cell layers thick. Obturator absent. Parietal tissue (two to) five cell layers thick. Nucellar cap present (sometimes poorly developed). Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear (heavily reduced). Endosperm haustoria? Embryogenesis?

Fruit A usually one-seeded (rarely two- och three-seeded) loculicidal capsule, with strongly accrescent calyx and persistent columella.

Seeds Aril absent. Seed coat testal. Testa multiplicative, vascularized (prominent postchalazal vascular bundles reaching to micropyle). Exotestal cells palisade, with thickened walls. Mesotesta aerenchymatous. Endotesta mainly collapsed. Tegmen collapsed. Perisperm not developed. Endosperm sparse or absent. Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, thick, fleshy, with simmondsin (a nitrile glycoside), aleurone grains and a liquid wax. Germination phanerocotylar?

Cytology n = 13

DNA Intron present in the plastid gene rpl2.

Phytochemistry Foliar flavonoids, condensed tannins, proanthocyanidins, anthocyanin, pinitol and simmondsinoid compounds present (simmondsin in jojoba-meal from ground seeds). Waxes consisting of high-molecular esters (C36 to C46 long straight-chain wax esters) of monoethylenic acids. Ellagic acid and alkaloids not found.

Use Oils and waxes (jojoba), medicinal plant.

Nakai in J. Jap. Bot. 18: 108. 10 Mar 1942

Stegnospermatineae Nakai in J. Jap. Bot. 18: 108. 10 Mar 1942

Genera/species 1/4

Distribution Coastal areas in northwestern and western Mexico and Central America, the West Indies.

Fossils Unknown.

Habit Bisexual, evergreen shrubs or small trees, more or less climbing.

Vegetative anatomy Phellogen in periderm. Secondary lateral growth anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements tracheids with bordered pits. Wood rays usually multiseriate (sometimes uniseriate), homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, or absent. Tyloses sometimes abundant. Phloem fibres absent. Sieve tube plastids P3c’f type, with a central polygonal protein crystal and a subperipheral dense ring of protein filaments. Nodes? Parenchyma, cortex and medulla with idioblasts containing sphaerite crystals.

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, entire, fleshy, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate, brochidodromous. Stomata anomocytic. Cuticular waxes? Lamina with idioblasts containing sphaerite crystals. Leaf margin entire, transparent.

Inflorescence Terminal or axillary, thyrsoid, raceme- or umbel-like.

Flowers Actinomorphic. Hypogyny. Tepals five, with quincuncial aestivation, sepaloid, persistent, free. ‘Petals’ (two to) five, with imbricate aestivation, probably staminodial in origin, free. Nectaries present in depressions at carpel bases. Disc absent.

Androecium Stamens five or eight to ten, in one whorl. Filaments connate at base, antepetalous stamens adnate to ‘petals’ (epipetalous). Anthers basifixed to subbasifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia probably present, petaloid.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, punctate, spinulate. Endexine massive.

Gynoecium Pistil composed of two to five connate alternisepalous (alternitepalous) carpels. Ovary superior, ab initio bilocular to quinquelocular, later unilocular. Style single, simple, short, or absent. Stigmas two to five, subulate, recurved, ventrally papillate, type? Pistillodium absent.

Ovules Placentation basal, gradually free central. Ovule one per carpel, amphitropous, erect, epitropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue. Nucellar cap absent. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A coriaceous capsule, dehiscing by three to five valves.

Seeds Aril present, covering larger part of seed. Seed coat exotestal-endotegmic? Exotesta palisade, with non-lignified cell walls. Endotesta? Exotegmen? Endotegmen enlarged, persistent. Perisperm copious, nutritious. Endosperm almost absent. Embryo curved, chlorophyll? Cotyledons two. Germination?

Cytology n = 36

DNA

Phytochemistry Very insufficiently known. Betalains (betacyanins, betaxanthins) present. Triterpene saponins probably abundant. Alkaloids? Pinitol? Proanthocyanidins not found.

Use Soap.

Doweld, Tent. Syst. Plant. Vasc.: xlii. 23 Dec 2001

Genera/species 2/c 15

Distribution Africa, Madagascar, western North America to South America, the West Indies, with their largest diversity in Mexico.

Fossils Unknown.

Habit Bisexual, usually small shrubs (sometimes perennial herbs, suffrutices or small trees?; ‘Talinella’ lianoid shrubs) with somewhat succulent leaves. Subterraneous organs often tuberous. Aerial parts often ephemeral.

Vegetative anatomy C₃ photosynthesis alternating with CAM photosynthesis. Stem epidermis usually with parallelocytic stomata, parallel to stem axis. Phellogen ab initio usually epidermal (rarely outer-cortical). Precocious or delayed initiation of stem periderm. Vessel elements with simple perforation plates; lateral pits alternate to pseudoscalariform or scalariform. Imperforate tracheary xylem elements usually libriform fibres (occasionally fibre-tracheids) with simple pits, septate. Wood rays multiseriate or absent. Axial parenchyma scanty vasicentric. Tyloses sometimes present. Thick-walled pericyclic extraxylary phloem fibre caps present. Sieve tube plastids P3cf type? Nodes 1:1, unilacunar with one leaf trace? Sclereids (thin-walled, elongated with lignified walls) often present. Mucilage idioblasts and tanniniferous cells abundant. Phloem parenchyma cells with phytoferritin? Epidermis and parenchyma often with numerous crystals or druses of calciumoxalate.

Trichomes Hairs multicellular, uniseriate, or absent; stem epidermis often with papillae or multicellular hairs.

Leaves Alternate (spiral), simple, entire, succulent, with ? ptyxis. Stipules and leaf sheath absent. Leaves usually with single or paired axillary scale-like vascularized prophylls representing rudimentary axillary short shoots (sometimes with tannins). Petiole vascular bundle transection arcuate; wing bundles present. Venation pinnate, brochidodromous? Stomata parallelocytic, transversely orientated. Cuticular wax crystalloids? Mesophyll with mucilaginous idioblasts. Leaf margin entire.

Inflorescence Terminal, thyrsoid paniculate cymose (sometimes raceme-like; in ‘Talinella’ congested heads), or flowers solitary axillary. Sepaloid floral prophylls (bracteoles) two, deciduous or persistent, often scale-like.

Flowers Actinomorphic. Hypogyny. Tepals (two to) five, with quincuncial imbricate aestivation, petaloid, caducous, free or connate at base. Nectaries extrastaminal, inserted at tepal bases. Disc absent.

Androecium Stamens c. 15 to c. 35. Filaments free from each other, free or adnate at base to tepals. Anthers dorsifixed, versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains polyforate or polyrugate, shed as monads, tricellular? at dispersal. Exine tectate, with columellate infratectum, spinulate.

Gynoecium Pistil composed of three (to five) connate carpels. Ovary superior, usually unilocular (in ‘Talinella’ multilocular). Style single, thin. Stigma bilobate or trilobate, type? Pistillodium absent.

Ovules Placentation free central. Ovaries numerous per ovary, anatropous?, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Apical cells of megasporangium radially elongate? Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit Usually a loculicidal valvicidal capsule (dehiscent from apex and/or base), often with caducous exocarp separating from fibrous persistent endocarp (capsule in ‘Talinella’ berry-like, juicy and mucilaginous; capsule in Amphipetalum a pyxidium, opening by a circumscissile operculum). Pericarp epidermis papillate. Capsule covered by dry remnants of perianth, stamens and style, these shed together as a calyptra.

Seeds Testa with strophiole near base. Seed coat? Exotesta? Endotesta? Tegmen? Perisperm copious, starchy? Endosperm absent? Embryo curved and sourrounding perisperm. Cotyledons two. Germination phanerocotylar?

Cytology n = 8 eller 12?

DNA 6 bp deletion in plastid gene ndhF.

Phytochemistry Virtually unknown. Betalains present.

Use Ornamental plants.

Link, Enum. Hort. Berol. Alt. 1: 291. 16 Mar-30 Jun 1821, nom. cons.

Tamaricales Bercht. et J. Presl, Přir. Rostlin: 233. Jan-Apr 1820 [‘Tamariscinae’]; Reaumuriaceae Ehrenb. ex Lindl., Intr. Nat. Syst. Bot.: 48. 27 Sep 1830 [‘Reaumurieae’]; Reaumuriales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 61. Sep-Oct 1835 [‘Reaumurieae’]; Tamaricineae Engl., Syllabus, ed. 2: 151. Mai 1898

Genera/species 3–4/75–90

Distribution Mostly drier regions of northern, northeastern and southwestern Africa, and Eurasia eastwards to East Asia, with their largest diversity in the Mediterranean and the irano-turanian areas; naturalized in North America.

Fossils Fossil wood of Tamarix has been found in Pleistocene (possibly also Pliocene) layers in North Africa.

Habit Usually bisexual (in Tamarix rarely dioecious), evergreen trees, shrubs or suffrutices. Xerophytes, rheophytes or halophytes.

Vegetative anatomy Phellogen ab initio usually superficially (rarely deeply) sited. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements usually libriform fibres (sometimes tracheids) with simple or (reduced) bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma paratracheal scanty, confluent, vasicentric, or banded. Vessel elements and/or parenchyma partially storied. Sieve tube plastids S type. Nodes? Heartwood often with resins etc. Calciumsulphate crystals sometimes present. Prismatic calciumoxalate crystals often abundant.

Trichomes Hairs unicellular or multicellular, uniseriate; salt glands common.

Leaves Alternate (spiral), simple, entire, usually small and scale-like or subulate (in Reaumuria coriaceous or fleshy, not scale-like), with ? ptyxis. Stipules and leaf sheath absent. Venation? (often single-veined). Stomata usually anomocytic (sometimes paracytic), often on adaxial side only, often deeply sunken. Cuticular wax crystalloids as tubuli or curled rodlets. Epidermis often with sunken, multicellular, salt-excreting glands. Leaf margin entire.

Inflorescence Usually terminal, simple or branched, catkin-, spike- or raceme-like, botryoid to paniculate (flowers in Reaumuria solitary terminal). Floral prophylls (bracteoles) absent.

Flowers Actinomorphic, usually small. Hypogyny. Sepals four or five (or six), with imbricate aestivation, persistent, usually free (in Reaumuria connate below). Petals four or five (or six), with usually imbricate (in Reaumuria contorted) aestivation, persistent or caducous, usually without scale-like appendage (in Reaumuria two adaxial scale-like appendages at petal base, without nectary), free or slightly connate at base, usually inserted at fleshy nectariferous disc. Nectariferous disc usually present below petals and stamens, often annular (rarely intrastaminal and/or extrastaminal, in Reaumuria absent).

Androecium Stamens in Myricaria 5+5 (antesepalous longer than antepetalous), in Tamarix and Myrtama four or five (to 14), in Reaumuria five to more than twelve (from ten primordia). Filaments usually connate at base, usually in a single group (in Reaumuria often five antepetalous groups), sometimes free, usually inserted on fleshy nectariferous disc, free from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, latrorse or extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tri- or tetracolpate, shed as monads or tetrads, bicellular (Tamarix) or tricellular (Myricaria germanica) at dispersal. Exine semitectate, with columellate infratectum, reticulate or microreticulate.

Gynoecium Pistil composed of (two or) three or four (or five) antepetalous connate paracarpous carpels; median carpel abaxial? Ovary superior, unilocular (sometimes seemingly bilocular to quinquelocular due to intrusive placentae). Stylodia (two or) three or four (or five), free or partially connate (absent in Myricaria). Stigmas with usually widened lobes, non-papillate, Wet type. Pistillodium absent?

Ovules Placentation parietal, basal or basal-parietal. Ovules two to numerous per carpel, anatropous, ascending, bitegmic, weakly crassinucellar. Micropyle endostomal. Outer integument two or three cell layers thick. Inner integument two or three cell layers thick. Megasporangium very thin. Parietal tissue approx. one cell layer thick (parietal cell often not dividing). Megagametophyte tetrasporous, Fritillaria, Adoxa, Drusa or Chrysanthemum type (also within the same species of Tamarix), sometimes 16-nucleate, bipolar. Synergids occasionally with a filiform apparatus. Endosperm development ab initio nuclear or cellular. Endosperm haustoria absent. Embryogenesis solanad. Polyembryony frequent.

Fruit A loculicidal capsule.

Seeds Seeds with long unicellular hairs (Reaumuria) or a chalazal coma of long hairs (Myricaria, Tamarix, Myrtama). Aril absent. Seed coat exotestal. Exotestal cells periclinally elongate, thick-walled. Endotestal cells thin-walled, with crystals. Tegmen entirely or largely absent. Perisperm often copious, thin. Endosperm usually absent (occasionally sparse then usually oily and proteinaceous; in Reaumuria sparse, starchy). Embryo large, straight, well differentiated, chlorophyll? Suspensor massive. Cotyledons two. Germination phanerocotylar.

Cytology n = (10) 12 (18, 24, 36) (Tamarix); n = 12 (Myricaria); n = 11 (Reaumuria)

DNA Intron present in plastid gene rpl2?

Phytochemistry Flavonol sulphates and bisulphates (kaempferol, quercetin or tamarixin), cyanidin, methylated and non-methylated ellagic acids, gallic acid, gallic and ellagic tannins, alkaloids, pinitol, and syringaresinol present. Myricetin, saponins and cyanogenic compounds not found.

Use Ornamental plants, medicinal plants, dyeing substances, tanning.

Cladogram of Tamaricaceae based on DNA sequence data (Gaskin & al. 2004).

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