MAGNOLIOPSIDA Brongn.

Brongniart, Enum. Plant. Mus. Paris: xxvi, 95. 12 Aug 1843 [’Magnolineae’]


Maximum likelihood (ML) majority-rule consensus tree of Magnoliopsida (Pan-Angiospermae) based on DNA sequence data (Soltis & al. 2011, slightly modified). Amborella, Nymphaeales and Schisandrales being successive sister-groups to all other angiosperms had more than 80% bootstrap (BS) support in the 17-gene analysis by Soltis & al. (2011). The position of Amborella as sister to all other angiosperms is highly supported by complete plastid genome sequence analyses (Kim, Yoo & al. 2004; Moore & al. 2007; Soltis & al. 2011; etc.). The sister-group relationship [Chloranthaceae+Magnoliidae] had a BS support of 85%. The clades [Magnoliales+Laurales] and [Canellales+Piperales] each had a BS support of 100%. The clade comprising Liliidae, Ceratophyllum and Tricolpatae was supported by 86%, whereas the support of Ceratophyllum as sister to Tricolpatae was only 68%. The BS support for Sabiaceae as sister to the remaining Tricolpatae (a trichotomy in this tree) was likewise relatively low (59%). Sabiaceae are sometimes recovered as sister to Proteales, yet with weak to moderate support (Qiu & al. 2006; Moore & al. 2008; Burleigh & al. 2009; Moore & al. 2011; Soltis & al. 2011). Gunnerales were sister to the Pentapetalae with a BS support of 99%. Ranunculales were as usual sister to the remaining Tricolpatae (BS support 100%). The positions of Didymelales and Trochodendrales as successive sister-groups to Gunneridae had a BS support of 98% and 100%, respectively (Moore & al. 2010; Soltis & al. 2011). Gunnerales were sister to the remaining angiosperms, the Pentapetalae, with a support of 100%. Superasteridae (Santalales to Asteridae) and Superrosidae (Saxifragales and Rosidae) were supported by 87% and 100%, respectively. In some other analyses Berberidopsidales were sister to Asteridae, and Caryophyllales were sister to these two groups. The position of Dilleniaceae as sister to Superasteridae was supported by 97% in Soltis & al. (2011). In some other analyses they were recovered as sister to, i.a., the remaining Pentapetalae, or to the clade [Superasteridae+Superrosidae] or to Superrosidae, although with fairly low support (Moore & al. 2010). The maximum parsimony (MP) consensus tree was largely identical to the ML tree, although Ceratophyllum was sister to Liliidae and Dilleniaceae were sister to Caryophyllales. – Extant angiosperms began to diversify in the mid-Jurassic, c. 170 Mya, according to unconstrained penalized likelihood analyses (Moore & al. 2007), and the five major mesangiosperm lineages diversified fairly rapidly during the earliest Cretaceous. The initial divergence of these five lineages was dated to 143.8+4.8 Mya, and the divergence of Chloranthus and Magnoliidae was dated to 140.3+4.8 Mya. The origins of the extant crown groups of Magnoliidae, Liliidae, and Tricolpatae were dated to branching points somewhat later in the Cretaceous: 130.1+4.4 Mya for Magnoliidae, 128.9+4.9 Mya for Liliidae, and 124.8+6.3 Mya for Tricolpatae. Divergence times and standard errors (in parentheses) in Mya for deep-level angiosperm nodes as estimated by penalized likelihood analyses were as follows: angiosperms 169.7 (3.46) – Nymphaeales+Illicium+Mesangiospermae 163.5 (2.63) – Illicium+Mesangiospermae 154.8 (2.53) – Mesangiospermae 143.9 (2.67) – Chloranthus+Magnoliidae 140.4 (2.54) – Magnoliidae 130.3 (2.20) – Liliidae+Ceratophyllum+Tricolpatae 143.1 (3.18) – Liliidae 129.1 (2.69) – Ceratophyllum+Tricolpatae 141.4 (2.97) – Tricolpatae 124.9 (3.43).

Pan-Angiospermae P. D. Cantino et M. J. Donoghue in Taxon 56, E23. 2007

Habit Bisexual or unisexual, trees, shrubs, lianas, suffrutices, or perennial, biennial or annual herbs.

Root Main root usually developing from radicula and usually as tap root with lateral roots (radicula sometimes ephemeral, all roots of mature plant being adventitious). Root cap and epidermis usually with common ontogenetic origin. Apical meristem open or intermediate. Vascular tissue diarch to pentarch. Lateral roots arising opposite or when diarch immediately adjacent to xylem poles. Phellogen deeply seated. Trichoblasts (differentiated cells forming root hairs) absent. Epidermis probably initiated from inner root cap layer.

Stem Shoot apex with tunica-corpus construction. Tunica two-layered. Phellogen usually superficial (initiated at or immediately below epidermis; sometimes deeply seated, i.e. initiated deep in cortex, inside pericycle or in phloem). Vascular bundles usually arranged in cylinder (annular in cross-section; sometimes in two or more concentric cylinders or as scattered bundles). Circular bordered pits when present without margo and torus. Perforation plates (tracheid:tracheid plates) with primarily scalariform pitting (secondarily simple). Wood fibres (imperforate tracheary xylem elements) and axial (wood) parenchyma usually present. Reaction wood with gelatinous fibres. Starch grains simple. Primary cell walls usually with pectic polysaccharides (mannans sparse). Cytoplasm not occluding sieve plate pores, containing P proteins. Sieve tube with sieve plate. Sieve tubes eunucleate (nucleated companion cells, non-nucleated sieve tube and P proteins developing from common mother cell). Albuminous Strassburger cells, functionally associated with sieve cells, not developing from same mother cell. Sieve tube plastids containing only starch grains (S type), or with protein inclusions as well (P type) (rarely with neither starch nor protein, S0 type). Nodes usually unilacunar or trilacunar (sometimes bilacunar or multilacunar) with usually one or three (sometimes two or more than three) leaf traces. Prophylls (including bracteoles) usually single or paired.

Leaves Leaf usually with petiole and lamina (lamina often sessile or leaf only consisting of petiole); lamina developing from primordial leaf apex. Leaf with acropetal development of venation. Stipules present or absent. Leaf sheath present (petiole base sheathing stem) or absent. Secondary venation pinnate, palmate or parallel; fine venation usually reticulate. Vein endings usually free (with open venation). Stomata often paracytic, with ends of guard cells level with pore; outer stomatal ledges producing vestibule. Cuticular wax crystalloids as parallel oriented platelets (Convallaria type). Leaf margin entire or serrate (with teeth). Leaf axil usually with bud (axillary bud).

Flower Flower bisexual or unisexual. Symmetry actinomorphic, bisymmetrical, zygomorphic or asymmetrical. Pedicel present or absent, usually provided with one or two floral prophylls (bracteoles). Floral parts spirally arranged or whorled in one or more series. Floral parts usually present in stable (sometimes unstable) numbers. Perianth differentiated into sepals and petals or undifferentiated. Tepals usually centripetally developing, with usually one or three traces, free or more or less connate, persistent or caducous. Outer tepals (sepals) enclosing or not enclosing remainder of floral bud. Floral parts when whorled usually trimerous, tetramerous or pentamerous. Floral nectaries of various origin present or absent.

Androecium Stamens usually centripetally (sometimes centrifugally) developing. Each stamen usually supported by one trace. Stamen usually differentiated into filament and anther (microsporangia sometimes embedded in distal part of stamen). Filament band-shaped or terete, usually narrow (sometimes wide and stout). Anther usually dithecal (microsporangia organized in two groups each with two sporangia; sometimes monothecal etc.), tetrasporangiate (sometimes disporangiate or polysporangiate), introrse, latrorse or extrorse. Microsporangia with at least outer secondary parietal cells dividing. Thecae usually dehiscing longitudinally (longicidally) by action of hypodermal endothecium (sometimes poricidally, valvicidally etc.). Endothecial cells usually elongated at right angles to longitudinal axis of anther. Tapetum usually secretory (glandular), with binucleate (sometimes uninucleate or multinucleate) cells (sometimes amoeboid-periplasmodial).

Pollen grains Microsporogenesis usually simultaneous (sometimes successive). Pollen grains usually mono- or triaperturate (sometimes other numbers), bicellular or tricellular at dispersal. Male gametophyte tricellular. Tectum continuous, discontinuous or absent. Infratectal layer usually columellate (sometimes granular or intermediate). Endexine usually thin, compact and only in apertural regions lamellate. Male gametangia (antheridia) absent. Male gametes two, without cell walls, without flagellae. Pollen tube with pectic outer cell wall, inner wall of callose, and with posterior callose plug and usually moderately fast growth (between c. 100 µm and c. 600 µm per hour). Siphonogamy prevailing. Pollen tubes growing inside secretions from stigma and through stylar canal and ovary locule, or through compitum (non-destructive pollen tube growing between megasporangial cells), callose being secreted onto cell walls behind growing pollen tube apex and over entire substigmatic region. Pollen tube usually penetrating micropylar end of ovule (porogamy) and proceeding into synergids (sometimes chalazogamy).

Gynoecium Carpels usually several or numerous, usually more or less connate (sometimes free). Carpels plicate or ascidiate, postgenitally usually fused (sometimes occluded by secretion). Compitum usually present. Carpellary distal-adaxial pollen-receptive part usually papillate or non-papillate, Dry or Wet (secretory) type stigma (receptive area sometimes present along free carpellary margins). Pollen grains deposited on receptive surface. Stigmatic surface also aiding in development of pollen tubes. Stylar part of gynoecium present or absent (stigma sessile), often hollow (with central canal).

Ovule Placentation of different types (axile, parietal, laminar etc.). Ovules usually anatropous, bitegmic or unitegmic and crassinucellar or tenuinucellar. Micropyle endostomal, bistomal or exostomal. Each integument one or several cell layers thick. Inner integument dermal or subdermal in origin. Parietal tissue present (ovule crassinucellar) or absent (ovule tenuinucellar). Megasporocyte usually single (archespore sometimes multicellular), hypodermal, present in centre of ovule. Cytoplasmically dense zone developing between megasporocyte nucleus in centre of cell and chalazal cell wall, this dense zone persisting through megasporogenesis. Megasporogenesis resulting in usually linear tetrad. Functional megaspore without sporopollenin and cuticle and usually chalazal. Megagametophyte usually monosporic and developing from chalazal cell. Meiosis I resulting in a dyad of two uninucleate cells divided by transverse cell wall. Meiosis II yielding a usually linear tetrad of megaspore cells, of which chalazal cell becomes megaspore and remaining three megaspores degenerating prior to initiation of megagametogenesis. Large central nucleus of functional megaspore surrounded by vacuole. Two nuclei migrating to opposite poles of cell after first mitosis. Two nuclei present at each pole after second mitosis, nuclei at micropylar pole becoming fusiform. Four nuclei present at each pole after third mitosis, cytokinesis now taking place. Resulting megagametophyte 8-nucleate and septacellular. Three cells at micropylar end becoming egg apparatus consisting of usually two synergids and one egg cell. Each synergid often containing a cuneate filiform apparatus, usually a small number of plastids and in general with thicker wall than egg cell. Three cells at chalazal end becoming antipodal cells, usually containing numerous plastids, and in many cases proliferating (dividing). Nuclei of these cells more or less fusiform (torpedo-shaped) and each with distinct nucleolus. Remaining two nuclei, polar nuclei, of seventh cell, central cell, moving to centre of megagametophyte. Here they are situated in a parietal band of cytoplasm. Female gametangia (archegonia) absent. Fertilization taking place some time between one day and one year following pollination. Fertilization double, i.e. one male gamete fusing with egg cell and second male gamete with central cell containing two free or fused polar nuclei (polar nuclei fusing in centre of central cell either immediately prior to or during fertilization process and subsequently moving into position adjacent to egg cell with its terminal nucleus). Triploid cell resulting from fertilization of diploid central cell usually developing into endosperm. Endosperm development cellular, helobial or nuclear. Endosperm nuclei triploid.

Fruit Fruit developing from ovary. Additional tissues often contributing to structures surrounding ovary resulting in pseudofruit. Fruit wall, pericarp, usually consisting of innermost endocarp, mesocarp outside this, and outer exocarp.

Seeds Seed coat consisting of one or several layers developed from integument(s). Endosperm sparse to copious, oily and/or proteinaceous and/or starchy (occasionally absent). Embryo ab initio cellular. Cotyledons usually two (sometimes one, rarely three or four), each usually provided with three vascular bundles (when one cotyledon, then usually with two main vascular bundles). Plumule usually terminal. Seedling with sympodial growth. Germination usually phanerocotyl (sometimes cryptocotyl).

Cytology and genes A genes are found only in angiosperms. A duplication event took place in the ancestor of angiosperms, which resulted in a D orthologue, usually expressed only in the ovules, and a C orthologue, only occasionally expressed in ovules, but in carpels and stamens as well.

Telomeres of Arabidopsis type (TTTAGGG)n. Entire nuclear genome duplicated. Second duplication of nuclear genes PEBP resulting in FT-like and TFL1-like gene families (first duplication leading to two PEBP gene families, MFT-like and FT/TFL1-like, and probably coinciding with evolution of seed plants).

Nuclear genes LEAFY and RPB2 present in single copy. knox genes extensively duplicated (A1 to A4). Nuclear genes AP1/FUL, paleoAP3 and PI (paralogous B-class genes) present, and possessing “DEAER” motif. Three copies of nuclear gene PHY (PHYBc(PHYA/PHYC)) and gene pair SEP3/LOFSEP duplicated (gene PHYA involved in, i.a., germination and etiolation response of seedling).

Nuclear gene euAP3 in angiosperms consisting of duplicated copy of gene paleoAP3 with 8 bp insertion causing a frame-shift mutation. APETALA3 and similar nuclear genes and PISTILLATA are paralogous B-class genes. B-function MADS-box genes are extremely important for the floral development.

The exon 5’ in the PI-homologues has a size of 42 base pairs in the basalmost clades Amborella and Nymphaeales. Illicium (Schisandrales) and all younger clades has a deletion of 12 bp in this exon 5’ resulting in a length of 30 bp.

Plastid gene ndhB extended 21 codons at 5’ end.

Phytochemistry Lignans, quercetin and/or kaempferol, O-methyl flavonols, dihydroflavonols, oleanane (triterpenoid), non-hydrolyzable tannins, tyrosine-derived cyanogenic compounds, apigenin and/or luteolin present. Lignins often derived from coniferyl and sinapyl alcohols, containing syringaldehyde (in positive Mäule reaction, syringyl:guaiacyl ratio higher than 2,0–2,5:1). Hemicelluloses often present as amyloid (xyloglucans).

Early angiosperm fossils of unknown affinity (adopted from Friis & al. 2011)

- Fossilized angiosperm leaves occur in Aptian and younger layers. A peculiar leaf fossil is the Aptian to Albian Trifurcatia flabellata, which is connected to the fossil genus Klitzschophyllites. These leaves are thick, circular in outline and with serrate margins. The venation is flabellate with more than 20 primary and secondary veins terminating at or between the leaf teeth. Gland-like structures are present along the leaf margins.

- Small male flowers from the Late Barremian to the Aptian (Early Cretaceous) of Portugal, with ten to 15 stamens having wide flat filaments and lateral thecae and trichotomocolpate pollen grains, are in some way similar to Amborella trichopoda, although the anthers were extrorse instead of introrse and the pollen surface was verrucate-rugulate.

- The Early Cretaceous (Early to Late Aptian, the Liaoning Province in northeastern China) probably limnic species of Archaefructus were herbaceous with multiple times divided leaves and leaflets. They had naked unisexual terminal reproductive organs usually arising in pairs, with female and male parts separate on the elongating axis. Bracts and perianth seem to have been absent. The seed-bearing carpel-like organs (possibly developing into follicles) were elongated and slightly stipitate and the stamen-like organs were present two to four together on a common stipe. The reproductive organ has been interpreted as a single bisexual flower or as an inflorescence consisting of numerous unisexual units. Archaefructaceae were recovered as sister to all other angiosperms in a combined morphological and 3-gene-analysis by Sun & al. (2002). Archaefructaceae comprise the two species Archaefructus liaoningensis Sun, Dilcher, Ji & Zhou and A. sinensis Sun, Dilcher, Ji & Nixon.

- Cronquistiflora and Detrusandra from the Turonian (93,5–89 Mya) in New Jersey had numerous spiral carpels and other floral parts.

- Caloda delevoryana, from Late Albian to Early Cenomanian (mid-Cretaceous) strata of the central United States, is represented by elongate infructescences bearing alternately arranged lateral branches each ending in a receptacle with numerous free stipitate carpels.

- The floral fossil Carpestella lacunata, from the Early to Middle Albian (Early Cretaceous) of Virginia, may have been related to extant basalmost angiosperms. The fossil consists mainly of a syncarpous gynoecium with 13 carpels covered by spiral scars of detached perianth and androecium.

- Caspiocarpus paniculiger is represented by a single shoot from the mid-Albian of Kazakhstan, having opposite palmately veined leaves and paniculate reproductive axes bearing follicle-like fruits.

- Cretovarium japonicum from the Coniacian to Campanian (Late Cretaceous) of Hokkaido (Japan) has perianth-like structures surrounding a trilocular inferior ovary with axile placentation (two separate placentae in each locule).

- Hidakanthus shiinae and Protomonimia kasai-nakajhongii are multicarpellate fruit structures from the Coniacian to Santonian (Late Cretaceous) of Hokkaido (Japan). Hidakanthus has c. 55 sessile carpels with oil cells and Protomonimia c. 170 or stipitate carpels on a concave receptacle. The seeds of Protomonimia are exotestal.

- Lesqueria elocata represents elongate infructescences from the Late Albian to Early Cenomanian (mid-Cretaceous) of Kansas and northern Texas. The gynoecium was apocarpous and multicarpellate, and bore up to c. 150 helically arranged laminar appendages and up to c. 250 stalked follicles.

- Xingxueina heilongjiangensis is an Aptian spicate inflorescence from the Heilongjiang Province in northeastern China. The proposed floral units are situated in an elongated helix. The monocolpate pollen grains have a reticulate exine.

- Zlatkocarpus brnikensis and Z. pragensis resemble the Chloranthaceae. They have spicate inflorescences with helically arranged floral units containing a single carpel subtended by an adnate bract and probably developed into a berry with resin bodies (possibly ethereal oil cells) in the pericarp.

- A number of flowers have been described from Cretaceous strata in New Jersey and Portugal. Mabelia (Turonian, Late Cretaceous) and Nuhliantha (Late Santonian, Late Cretaceous) were trimerous and unisexual, with six tepals and three extrorse stamens containing monocolpate (monosulcate) or trichotomocolpate pollen grains. With the exception of a pistillodium in Nuhliantha no female organs are known. The floral morphology suggests a monocotyledonous affinity. Microvictoria (Turonian) consists of bisexual pedicellate flowers bearing numerous tepals (or bracts?), flattened staminodia and stamens and carpels. They are similar to flowers in Nymphaeaceae, although this relationship may be questioned.

- Numerous fossilized fruits and seeds occur in Cretaceous layers. The Early Cretaceous Anacostia represents small single-seeded berries with exotestal anatropous seeds. Trichotomocolpate pollen grains with reticulate tectum are often associated with these fruits. The flowers seem to have been apocarpous with several or many carpels. Couperites likewise comprises single-seeded berries with exotestal anatropous seeds, often found together with monocolpate pollen grains with reticulate sexine similar to the Clavatipollenites pollen type.

- Angiosperm pollen grains are frequent in Cretaceous beds. The Afropollis form genus includes spheroidal, acolumellate coarsely reticulate grains, usually with a granular infratectum. They may be zonacolpate, monocolpate or inaperturate and either isopolar or heteropolar. Afropollis occurs in Early Cretaceous layers from the Barremian to the Cenomanian. The angiospermous origin of Afropollis has been questioned, i.a. due to the presence of a thick laminar endexine unknown among extant flowering plants. The similar Schrankipollis form genus from the Early Cretacous represents zonacolpate, loosely reticulate pollen grains with columellate infratectum.

- Clavatipollenites comprises monocolpate pollen grains with a reticulate exine, finely verrucate colpus membrane and indistinct colpus margins. Many Clavatipollenites grains have been assigned to Chloranthaceae, and to Ascarina in particular, although the form genus Clavatipollenites certainly represents several different early angiosperm clades.

- Retimonocolpites dividuus is an Early Cenomanian monocolpate pollen with a reticulate exine and an aperture encircling most of the grain. Brenneripollis represents monocolpate pollen grains with irregularly reticulate exine and columellate infratectum, whereas the similar Pennipollis has acolumellate infratectum.

- Liliacidites is monocolpate or trichotomocolpate and has a graded reticulum with the small lumina concentrated in the equatorial area. This Early Cretaceous to Early Cenozoic type resembles pollen grains of extant monocotyledons. Similipollis, also with possible monocotyledonous affinities, has the small lumina of the reticulum concentrated in the polar area.

- The Barremian to Cenomanian Stellatopollis comprises monocolpate pollen grains with columellate infratectum and reticulate exine beset with clavate supratectal elements which are borne in a stellate pattern. The Albian Transitoripollis pollen type is monocolpate and has continuous verrucate to microechinate tectum and granular infratectum. The Barremian to Aptian Tucanopollis is similar to Transitoripollis, but the colpus is sometimes nearly circular in outline. The large Lethomasites pollen type is also monocolpate and tectate with granular infratectum, although the tectum is perforate.

Systematics The main clades of flowering plants are briefly presented below. The potential synapomorphies are mainly adopted from Peter F. Stevens, “The Angiosperm Phylogeny Website”, version 9 (June 2008, updated in July 2012). More comprehensive descriptions are given for Magnoliidae (the magnolids), Liliidae (the monocots), Asteridae (the asterids), and Rosidae (the rosids).

Nymphaeidae J. W. Walker ex Takht., Divers. Classif. Fl. Pl.: 74. 24 Apr 1997

[Nymphaeales+[Schisandrales+[[Chloranthaceae+Magnoliidae]+[Liliidae+[Ceratophyllum+Tricolpatae]]]]]

Potential synapomorphies: Vessels present in wood (xylem). Vessel elements with elongated scalariform perforation plates. Wood fibres (imperforate tracheary xylem elements) present. Axial parenchyma diffuse or diffuse-in-aggregates. Pollen grains monosulcate (anasulcate). Exine tectate, columellate, reticulate to perforate. “DEAER” motif of AP3 and PI genes absent (lost). – Nymphaeidae comprise all extant flowering plants except Amborella trichopoda. Wood characters in general are extremely variable and very much depend on the environmental conditions. The wood anatomical variation is also, naturally, correlated with size, age and life style of the plant. Hence, the presence of different types of perforation plates, pits, imperforate tracheary xylem elements, wood (axial) parenchyma, and wood rays only exceptionally present features useful at taxonomically higher levels.

Illiciidae C. Y. Wu in Acta Phytotaxon. Sin. 40: 291. 2002

[Schisandrales+[[Chloranthaceae+Magnoliidae]+[Liliidae+[Ceratophyllum+Tricolpatae]]]]

Potential synapomorphies: Vessels present in stem xylem. Ethereal oils in spherical idioblasts (oil cells, making leaves and tepals pellucid-punctate). Tension wood absent. Anther wall with dividing outer secondary parietal cell layer. Sexine reticulate. Infratectal layer columellate. Carpels plicate, sealed by postgenital fusion of their margins. Nucellar cap present. Deletion comprising 12 bp (representing four amino acids) in exon 5’ of nuclear gene PI. – Illiciidae include all angiosperms except Amborella and Nymphaeales.

Mesangiospermae M. J. Donoghue, J. A. Doyle & P. D. Cantino in Taxon 56, E23. Aug 2007

[[Chloranthaceae+Magnoliidae]+[Liliidae+[Ceratophyllum+Tricolpatae]]]

Potential synapomorphies: Endomycorrhiza vesicular-arbuscular. Outer epidermal walls of root elongation zone provided with cellulose fibrils transversely orientated in relation to root axis. Perianth trimerous. Tepals whorled, trimerous. Stamens whorled. Megagametophyte bipolar, septacellular, octanucleate. Antipodal cells persistent. Endosperm triploid. Benzylisoquinoline alkaloids and polyacetate-derived anthraquinones present. – Mesangiospermae embrace flowering plants other than the ANITA grade (i.e. Amborella, Nymphaeales and Schisandrales). It is questionable whether whorled floral parts is a synapomorphy at this level. Both spiral and whorled tepals, stamens and carpels are frequent in Magnoliidae.

[Chloranthaceae+[[Magnoliales+Laurales]+[Canellales+Piperales]]]

Potential synapomorphies: Seed coat endotestal. Sesquiterpenes present.

Magnoliidae Novák ex Takht., Sist. Filog. Cvetk. Rast.: 51. 4 Feb 1967 (magnolids)

[[Magnoliales+Laurales]+[Canellales+Piperales]]

Potential synapomorphies: Vessel elements solitary and in radial multiples. Sieve tube plastids often containing polygonal protein crystals. Leaf margin entire. Stamens numerous, spirally arranged. Anthers extrorse. Hypostase present. Nucellar cap present. Antipodal cells ephemeral. Raphal bundle branches present at chalaza. Galbacin and verguensin (lignans) present. Licarin (neolignan) sometimes present. Asarone (phenylpropane)?

Habit Usually woody (in Piperales usually herbaceous). Often aromatic.

Vegetative anatomy Medulla septate, with sclerenchymatous diaphragmata. Primary stem with eustele or (pseudo)siphonostele (sometimes atactostele), with separate (sometimes scattered) vascular bundles or with continuous vascular cylinder (rarely with several concentric cylinders). Secondary lateral growth rarely anomalous or absent (Piperales). Wood elements (and cambium) sometimes storied. Vessel elements with usually scalariform or simple (sometimes opposite or reticulate) perforation plates; lateral pits alternate, scalariform or opposite, simple or bordered pits; vessel elements sometimes absent and replaced by tracheids. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate, or absent. Secondary phloem often stratified. Sieve tube plastids usually Psc type (sometimes Ss, S0, Pcs, Psf, Pcsf, P2c, or Pc type). Secretory cavities with resins present or absent. Wood rays sometimes with oil cells or crystals. Idioblasts with ethereal oils often present at least in parenchyma. Sclereids often present. Mucilage ducts sometimes present. Silica bodies sometimes present. Calciumcarbonate as prismatic, rhomboidal or acicular crystals, styloids, druses or crystal sand sometimes present.

Trichomes Hairs unicellular or multicellular, usually uniseriate, simple or branched (stellate, dendritic, furcate, peltate, lepidote, candelabra-shaped, T-shaped or fimbriate), or absent; glandular hairs usually absent (pearl glands occasionally present).

Leaves Usually alternate (sometimes opposite, rarely verticillate), simple, usually entire (rarely lobed or scale-like), with conduplicate, supervolute, involute, convolute, curved or flate ptyxis. Stipules usually absent (sometimes intrapetiolar or ocreate and enclosing young leaf); leaf sheath absent (petiole rarely sheathing). Venation usually pinnate, eucraspedodromous or brochidodromous (rarely campylodromous, palmate, acrodromous or actinodromous, triplinerved or pedate). Stomata usually paracytic (sometimes tetracytic, rarely anomocytic, actinocytic, cyclocytic or helicocytic). Cuticular wax crystalloids as platelets (sometimes parallel), rodlets (often transversely ridged Aristolochia type crystalloids) or tubuli (sometimes as clustered tubuli of Berberis type), chemically characterized usually by presence of palmitone (hentriacontan-16-one) and absence of nonacosan-10-ol (nonacosan-10-ol present in Canellales). Lamina often gland-dotted. Domatia sometimes present in abaxial vein axils. Mesophyll and epidermis usually with idioblasts (secretory cavities) containing ethereal oils (sometimes calciumoxalate crystals, resin or mucilage) or sclereids, sometimes with calciumoxalate druses. Sclerenchymatous idioblasts with branched sclereids of various kinds (also asterosclereids) or fibres often present. Silica bodies sometimes present. Leaf margin usually entire (sometimes serrate with monimioid teeth).

Inflorescence Cymose panicle, umbellate, corymb, rhyrse, cyme or fasciculate (sometimes spadix; rarely capitate, spicate or raceme), or solitary. Floral prophyll (bracteole) often single, median, adaxial (sometimes absent).

Flowers Usually actinomorphic (in Piperales usually zygomorphic). Usually hypogyny (rarely half epigyny or epigyny), sometimes with urceolate, campanulate, cupular or infundibuliform receptacle surrounding floral parts. Tepals (two or) three (or 2+2) or 3+3(+3) (sometimes 2+2+2 or 4+4, rarely to more than 50), with valvate (usually outer) or imbricate (outer or inner; rarely decussate) aestivation, spiral or whorled, sepaloid (usually outer) or petaloid (usually inner), usually free or connate only at base (sometimes intirely connate); tepals sometimes absent. Nectary usually absent (sometimes with nectariferous disc, staminal nectariferous glands or adaxial nectaries inside perianth tube). Disc usually absent.

Androecium Stamens (one or) two to c. 20 to more than 200, laminar (foliaceous), spiral or whorled, not differentiated into filament and anther, with separate microsporangia embedded in distal part (adaxially, laterally, abaxially or apically), or differentiated into filament and anther. Filaments when present usually free from each other (sometimes partially or entirely connate; occasionally adnate to pistil into synandrium or gynostemium), usually free from tepals, sometimes with basal nectariferous glands. Anthers when present usually basifixed, non-versatile, usually free (rarely adnate to style), usually tetrasporangiate (sometimes disporangiate), sometimes with transversely septate thecae, extrorse, latrorse or introrse, longicidal (dehiscing by longitudinal slits) or valvicidal (dehiscing by valves), sometimes connate into synandrium; or microsporangia four, usually adaxial or lateral (sometimes abaxial), usually introrse or latrorse (sometimes extrorse), longicidal (dehiscing by longitudinal slits) or valvicidal (dehiscing by valves). Tapetum secretory or amoeboid-periplasmodial. Staminodia extrastaminal, intrastaminal, or absent.

Pollen grains Microsporogenesis simultaneous, successive or intermediate. Pollen grains usually monosulc(ul)ate (anasulc[ul]ate), monoporate or inaperturate (sometimes di- or trisulc[ul]ate, trichotomosulcate, polyporate, etc.), boat-shaped, usually shed as monads, usually bicellular at dispersal. Exine with granular, columellate or intermediary infratectum.

Gynoecium Carpels usually ten to more than 50 (sometimes one or few), spiral or whorled, free or more or less connate (sometimes paracarpous); carpel plicate to conduplicate (sometimes basally ascidiate and not differentiated into ovary and style), usually postgenitally incompletely or entirely occluded by fusion and/or secretion, with secretory canal, often open and filled by secretions, or without canal. Carpels often not differentiated into ovary, style and stigma. Ovary usually superior (sometimes inferior, rarely semi-inferior), unilocular to 20-locular (or more). Stylodium or style single, terminal, usually simple (occasionally lobate), or stylodia lateral to gynobasic, or absent (pollen tube transmitting tissue well developed). Stigma capitate or lobate, terminal or decurrent, papillate or non-papillate, Dry or Wet type. Nectar sometimes secreted from exposed carpel surfaces. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation parietal, laminar, marginal, submarginal, apical, subapical, basal, subbasal, lateral or axile. Ovules (one or) two to more than 100 per carpel (or one per ovary), usually anatropous (sometimes hemianatropous, orthotropous, hemiorthotropous or campylotropous), ascending, horizontal or pendulous, apotropous, usually bitegmic (sometimes unitegmic, rarely tritegmic), usually crassinucellar (sometimes tenuinucellar). Micropyle endostomal or bistomal (rarely exostomal), sometimes Z-shaped (zig-zag). Funicular obturator sometimes present. Archespore usually unicellular (rarely multicellular). Nucellar cap present or absent. Nucellar beak present or absent. Megagametophyte usually monosporic, Polygonum type (sometimes tetrasporic, Fritillaria or Peperomia type, etc.). Synergids usually with filiform apparatus. Antipodal cells ephemeral or persistent, sometimes proliferating. Endosperm development usually cellular (sometimes nuclear). Chalazal or micropylar endosperm haustoria sometimes present. Embryogenesis onagrad, asterad, piperad, or irregular.

Fruit A usually fleshy (sometimes leathery or more or less woody), dehiscent or indehiscent, apocarpous follicular fruit or a multifolliculus, or a dry or fleshy syncarp, a loculicidal (and occasionally septicidal) capsule, or a drupe (sometimes a single-seeded berry or an assemblage of achenes, berries, drupelets, dry follicles, samaras, etc.).

Seeds Perisperm usually not developed (in Piperales usually copious, starchy). Endosperm copious (to scarce), oily (occasionally also with compound starch grains), or absent. Embryo straight or slightly curved, more or less differentiated or undifferentiated, without chlorophyll. Cotyledons usually two.

DNA Deletion of 30 bp (corresponding to 10 amino acids) present in PI-derived motif in nuclear gene AP3 in most Magnoliales. Gene PI duplicated in Laurales. Intergenic inversion of c. 200 bp present in plastid inverted repeat in Laurales. Nuclear gene PHYE lost in Piperales.

Phytochemistry Flavonols (kaempferol, quercetin, etc.), 5-O-methylflavonols, flavones, flavanonols, diarylpropanes, catechins, cyanidin, monoterpenes, diterpenes (kauranes, clerodanes etc.), triterpenoids (tetracyclic etc.), oleanolic acid derivatives, sesquiterpenes, drimane sesquiterpenoids, sesquiterpene lactones, allyl- and propenylphenols, oxyphenols and other aromatic substances, phenylpropanes, caffeic acid, tannins, proanthocyanidins, aporphine alkaloids (aporphines, oxoaporphines, etc.), aporphine derivatives, benzylisoquinoline and other isoquinoline alkaloids, protoberberine alkaloids, C-methylated alkaloids, indole alkaloids, polyketide alkaloids (e.g. hallucinogenic pyridine alkaloids), quercetin glycosides, cyanogenic glycosides (dhurrin, triglochinin etc.), α-pyrones, myristicin, N-(cinnamoyl)-tryptamines, lignans (austrobailignan, veraguensin, dihydrocubebin), neolignans (aryltetralin, diaryltetrahydrofurans, etc.), lignoids, ethereal oils, phenanthrenes, aristolochic acids, polyketides, nitrophenyl ethan, germacrane-like compounds, myo-inisitol, syringaresinol, pinitol, kadsurin A, galbacin, licarin A, naphthoquinones, cinnamoylamides, arbutin, asarone, and amides present. Ellagic acid, gallo- and ellagitannins not found.

Fossils Examples of early fossils not assigned to any particular magnoliid clade are as follows.

- Araripia florifera, from Upper Aptian to Lower Albian of Brazil, is represented by a flowering shoot with decussate trilobate leaves. The tepals and bracts are spirally inserted on a cupular receptacle. It has not been possible to assign this fossil to a particular clade of the Magnoliidae.

- Detrusandra mystagoga, from the Turonian (Late Cretaceous) of New Jersey, is another uplaced magnoliid fossil comprising pedicellate flowers with cup-shaped receptacle, on which bracts and tepals are spirally inserted. The numerous stamens are situated on the inner upper part of the receptacle. The four microsporangia are adaxial on each stamen. The pollen grains are monocolpate with a reticulate exine. The five carpels are plicate and free and the stigmas are bilobate. Ovules are numerous and arranged in two ventral rows.

- Cronquistiflora sayrevillensis is a Turonian floral fossil from New Jersey. It resembles the above two fossils, having spirally arranged bracts and tepals, but the receptacular cup is shallow. The numerous free carpels are spirally inserted and terminating in a peltate stigma. The ovules are interpreted as orthotropous, bitegmic and with an endostomal micropyle.

- Canrightia resinifera, from the Aptian to Early Albian of Portugal, comprises fossilized flowers, fruits and seeds. The tepals are arranged in one whorl and connate, forming a hypanthium. The pollen grains are monocolpate with a reticulate exine and columellate infratectum. The two to five carpels are unilocular, connate and adnate to the tepals. The single ovule is orthotropous, endotestal-endotegmic and has a well-developed endothelium (also present in the extant Lactoris fernandeziana). Resin bodies are frequent on the ovary walls, indicating the presence of oil cells. The fruit was probably a berry.

Systematics Cuticular wax crystalloids as transversely ridged rodlets (Aristolochia type) are of high systematic significance characterizing Magnoliales, Laurales and Piperales. Sporadically, they also occur in various other taxa. Chemical analyses show that transversely ridged rodlets clearly differ in their composition. Waxes of one group are characterized by ketones, whereas a second group completely lacks ketones and is dominated by alkanes. Hentriacontan-16-one (palmitone) was found to be characteristic for transversely ridged rodlets in Aristolochia, Laurus, and Paeonia. Transversely ridged rodlets or related crystals grow from total waxes of all species but never crystallize from individual compounds such as alkanes or palmitone. Transversely ridged crystals are formed by self-assembly based on a slow crystallization process and the presence of additives.

[Magnoliales+Laurales]

Potential synapomorphies: Cuticular wax crystalloids as annularly ridged rodlets; main wax palmitol. Stamens whorled. Pollen grains with lamellate endexine. Carpel cross-zone initiated late. Placentation basal. Ovules one (or two) per carpel, orthotropous, apotropous. Fruitlets single-seeded.

[Canellales+Piperales]

Potential synapomorphies: Nodes 3:3. Carpels whorled. Flavonols and aporphine alkaloids present.

[Liliidae+[Ceratophyllum+Tricolpatae]]

Potential synapomorphies: Tepals present in two whorls (secondarily one whorl). Stamens present in two whorls (secondarily one whorl), outer whorl antesepalous, inner whorl antepetalous.

Liliidae J. H. Schaffner in Ohio Naturalist 11: 413. Dec 1911 (monocotyledons)

Habit Usually perennial herbs (sometimes secondarily woody). Growth basically sympodial (sometimes monopodial). Prophylls usually single, adaxial. Often with bulb or corm rich in polysaccharides. Usually without idioblasts containing ethereal oils etc.

Root Ectomycorrhiza usually absent (Arum type or Paris type arbuscular mycorrhiza occasionally present). Radicula usually ephemeral, early withering and replaced by adventitious roots from stem (or sometimes from hypocotyl). Root cap and root epidermis of different ontogenetic origin. Root epidermis developing from outer cortical layer. Inner epidermis absent. Tunica two-layered. Single- or multi-layered velamen often present. Trichoblast present in atrichoblast or trichoblast cell pair further from apical meristem. Trichoblasts (small densely staining cells giving rise to root hairs) present in vertical files with small proximal cell (near root apex) producing root hairs, or hypodermal cells (particularly those with velamen) sometimes dimorphic (root hairs pushing up through overlying cells). Phellogen usually absent (rarely superficial; phellogen initially developing immediately inside exodermis). Vascular root tissue oligarch to polyarch, usually medullated. Lateral roots arising opposite phloem poles. Endodermal cells with U-shaped wall thickenings. Secondary growth usually absent; anomalous when present. Vessels present only in roots. Vessel elements with scalariform or simple perforation plates. Tracheids absent.

Stem Distinct bark and medulla usually absent. Primary thickening meristem at least usually absent. Vascular system usually consisting of numerous separate scattered bundles (atactostele), often amphivasal, closed (sometimes consisting of one or two or more concentric cylinders of bundles). Interfascicular (vascular) cambium usually not developing (vestigial cambium occasionally present; vascular bundles sometimes with weakly developed cambial layer). Thickening of main axis sometimes taking place by division and enlargement of ground parenchyma cells (diffuse secondary growth) or by special type of cambium arising in parenchyma outside primary vascular system. Secondary lateral growth usually absent (anomalous when present). Amphivasal vascular bundles usually present. Vessel elements usually absent in stem and leaves; when present with usually scalariform (sometimes simple, rarely reticulate) perforation plates; lateral pits scalariform or alternate, simple or bordered pits. Imperforate tracheary xylem elements tracheids. Phloem parenchyma absent. Vascular bundles with late developing thick-walled sieve tubes without companion-cells. Sieve tube plastids usually P2c or P2cf types (sometimes P2cs or P2cfs type, rarely Ss type). Laticifers and latex sometimes present. Schizogenous ducts and cavities with resins and oils sometimes present. Secretory (often lysigenous) mucilage cavities and ducts often present. Tanniniferous idioblasts sometimes present. Idioblasts with suberised cell walls and containing aromatic oils and resins sometimes present. Silica bodies present or absent (special cells, isodiametric stegmata connected to fibres, with cap-shaped, druse-like, stellate, conical, boat-shaped or spherical silica bodies, sometimes present). Mucilage cells, ducts and chambers present, often with calciumoxalate as raphides, pseudo-raphides, styloids, crystal sand, or rhomboidal, cuboid or acicular crystals. Epidermal cells often with crystals.

Trichomes Hairs usually absent (sometimes unicellular or multicellular, uniseriate or multiseriate, simple or branched, T-shaped, stellate, peltate or lepidote, rarely prickly or dendritic); microhairs sometimes present; glandular hairs usually absent (multicellular glandular hairs or tricellular glandular microhairs rarely present).

Leaves Usually alternate (often tristichous; rarely opposite or verticillate), usually simple and entire (sometimes compound and/or lobed), often linear, usually bifacial, often subulate (sometimes equitant), with revolute, supervolute, involute, convolute, conduplicate, plicate, explicative, adplicate, curved or flat ptyxis, not differentiated into petiole and lamina (sometimes differentiated into pseudopetiole and pseudolamina, ptyxis then referring to pseudolamina); when seemingly present, then not homologous to petiole and lamina in other angiosperms (true lamina absent). Majority of leaf usually developing from hypophyll (pseudolamina developing from leaf base zone). Vorläuferspitze (precursor tip, abaxial unifacial conical or cylindrical protrusion at apex of mature leaf, representing upper part of leaf) often present. Stipules usually absent (rarely present as axillary scales inside leaf sheath); leaf sheath open or closed, often well developed, or absent (rarely with axillary intravaginal scales/colleters, squamulae intravaginales, in Alismatales). Ligule(s) (developing from adaxial intercalary meristems in transition zone between hypophyll and hyperphyll) sometimes present. Venation parallelodromous or pinnate-parallelodromous (rarely acrodromous, pedate, curvipalmate, campylodromous, reticulodromous, actinodromous or camptodromous), acropetally and basipetally developing from base, converging towards apex (closed at apex), or secondary pseudopinnate or pseudopalmate types; intermediate and other veins basipetal from apex; vein endings not free. Imperforate tracheary xylem elements tracheids. Stomata paracytic (in lines, parallel to long axis of leaf), brachyparacytic (cell divisions oblique), tetracytic, anomocytic, or tricytic (sometimes cyclocytic, hexacytic or polycytic); neighbouring cells with oblique or non-oblique divisions. Cuticular wax crystalloids as parallel platelets (Convallaria type), as longitudinally aggregated rodlets (Strelitzia type, chemically dominated by wax esters) or as unordered platelets, rodlets or filiform reticulate processes, or absent. Dimorphic hypodermal cells sometimes present. Epidermis often with bulliform cells or with idioblasts containing silica bodies. Mesophyll often with sclerenchymatous idioblasts, or with mucilaginous idioblasts containing calciumoxalate raphides, druses, prismatic or rhomboidal crystals, styloids, or crystal sand (sometimes with idioblasts containing ethereal oils). Schizogenous laticiferous cavities sometimes present. Tanniniferous cells sometimes abundant. Leaf margin usually entire (sometimes serrate or spinose-serrate). Leaf teeth non-glandular.

Inflorescence Simple or compound, cymose pseudumbel or pseudoraceme, spicate or capitate, panicle, fascicle, corymb, thyrsoid or spadix, often compound and consisting of bostrychoid or helicoid monochasial partial inflorescences, or raceme or spike, sometimes subtended by one or several spathae (enlarged bract usually surrounding spadix); or flowers solitary. Floral prophylls (bracteoles) usually single, usually adaxial, bicarinate (rarely lateral, pairwise or absent; axillary flowers sometimes possibly representing reduced lateral inflorescence branches).

Flowers Actinomorphic or zygomorphic, often with median outer tepal adaxial (rarely asymmetrical). Flowers usually pentacyclic and trimerous. Hypanthium rarely present. Hypogyny or epigyny (rarely half epigyny). Usually trimerous (rarely dimerous, tetramerous or pentamerous), usually pentacyclic. Perianth pseudomonocyclic (each providing sector for perianth tube when present). Tepals (2–)3(–7)+(2–)3(–7), in two whorls, often similar, with tepals of successive whorls alternating, with usually open (sometimes imbricate or valvate; rarely contorted, conduplicate, induplicate, etc.) aestivation, all sepaloid or petaloid, or outer tepals sepaloid and inner tepals petaloid, free or more or less connate into infundibuliform, tubular or urceolate perianth, or absent (tepals sometimes membranous or chartaceous, sometimes modified into scales, bristles or hairs); median outer tepal usually abaxial (rarely adaxial); each tepal provided with three leaf traces. Nectaries as septal nectaries, usually infralocular, or androecial nectaries (rarely tepal nectaries), or absent. Disc usually absent (nectariferous disc occasionally present).

Androecium Stamens (2–)3(–6)+(2–)3(–6) (sometimes one or three, rarely two, five, 6+3, 6+4 or up to more than 1.000), usually as many as tepals, usually antesepalous (sometimes antepetalous), whorled. Staminal primordia often associated and/or stamens vascularized from tepal trace. Anther and filament sharply distinguished. Filaments free from each other or more or less connate (rarely connate into synandrium), free from or adnate to tepals (epitepalous; rarely to style). Anthers usually dorsifixed, basifixed or subbasifixed (sometimes centrifixed), versatile or non-versatile, usually tetrasporangiate (rarely disporangiate or trisporangiate), introrse, latrorse or extrorse, usually longicidal (dehiscing by longitudinal slits; sometimes poricidal, dehiscing by one or two apical or subapical pores, or transverse slits). Endothecium developing from outer secondary parietal cell layer; inner secondary parietal cell layer dividing. Tapetum usually secretory (sometimes amoeboid-periplasmodial), with uninucleate to quadrinucleate cells. Staminodia present (rarely petaloid) or absent (female flowers often with staminodia).

Pollen grains Microsporogenesis usually successive (sometimes simultaneous); tetrads usually tetragonal. Pollen grains usually monosulcate (monocolpate; sometimes monosulcoidate, disulcate, trichotomosulcate, mono- to polyporate, spiraperturate, inaperturate, etc.), usually shed as monads (rarely dyads, tetrads, polyads or cryptotetrads), usually bicellular (sometimes tricellular) at dispersal. Exine usually with columellate (sometimes granular) infratectum. Endexine usually absent. Callose plugs of pollen tube usually irregularly spaced and incomplete.

Gynoecium Pistil composed of (one to) three (to more than 100) usually more or less connate (eusyncarpous or paracarpous; sometimes secondarily free), usually whorled (rarely seemingly spiral), antesepalous carpels; median carpel usually abaxial; carpel plicate or conduplicate; fusion congenital intercarpellary and/or postgenital; carpel ascidiate to plicate or intermediate, seemingly occluded by secretion. Ovary superior or inferior (rarely semi-inferior), (unilocular to) trilocular or multilocular. Style single, simple, with stylar canal (hollow), or stylodia terminal, lateral (rarely gynobasic), or absent. Stigma capitate, punctate, peltate or lobate (sometimes infundibuliform, copular, etc.), or stigmas linear, papillate or non-papillate, Dry or Wet type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation axile, parietal, basal, subbasal or apical (rarely laminar or marginal). Ovules one to more than 100 per carpel (rarely hundreds to tens of thousands or more), anatropous or campylotropous (sometimes semicampylotropous, hemianatropous, amphitropous, orthotropous, pleurotropous, plagiotropous, etc.), ascending, horizontal or pendulous, apotropous or epitropous, usually bitegmic (sometimes unitegmic, rarely ategmic), crassinucellar or tenuinucellar (sometimes pseudocrassinucellar or pseudotenuinucellar). Micropyle endostomal or bistomal (sometimes exostomal). Funicular obturator sometimes present. Parietal cell formed from archesporial cell (parietal tissue usually one cell layer thick) or absent. Periclinal cell divisions sometimes taking place in megasporangial tissue (parietal cell then not formed). Nucellar cap sometimes formed by periclinal divisions of megasporangial epidermis. Epidermal cells of megasporangium sometimes forming ‘nucellar pad’. Megagametophyte usually monosporic, Polygonum type (occasionally Oenothera type, or disporic, Allium, Veratrum lobelianum, Endymion, Scilla types, or tetrasporic, Adoxa, Fritillaria or Drusa types). Synergids often with filiform apparatus. Antipodal cells usually persistent, sometimes proliferating. Endosperm development usually helobial (sometimes nuclear, rarely cellular), with distinct (large) micropylar and (small) chalazal chambers usually developed. Endosperm haustoria chalazal and/or micropylar or absent. Embryogenesis asterad or onagrad (sometimes caryophyllad, chenopodiad, or solanad).

Fruit Usually a loculicidal capsule (rarely septicidal, septifragal, poricidal, ventricidal, irregularly dehiscent or indehiscent), a berry, nut or drupe (sometimes a nut-like caryopsis, follicle, samara or schizocarp, or an assemblage of achenes, drupelets or berrylets or a multifolliculus).

Seeds Exotesta usually with thin phytomelan layer on epidermal cell walls. Perisperm usually not developed (sometimes well developed, with lipids and proteins or compound starch grains). Endosperm copious to sparse, often with starch (with simple or compound starch grains), and/or lipids, aleurone and hemicellulose, or absent. Chalazosperm developed or absent. Embryo straight to curved, well or poorly differentiated, sometimes covered with discoid or conical embryostega, testal operculum and surrounded by micropylar collar, with or without chlorophyll. Cotyledon one (rarely with rudimentary additional cotyledon), terminal, sometimes photosynthesizing, with closed sheath, usually unifacial (hyperphyllar; sometimes bifacial), assimilating and haustorial, usually with two main vascular bundles. Plumule lateral. Cotyledon hyperphyll elongate or compact, dorsiventrally flattened, assimilating or not assimilating, sometimes modified into haustorium or nutrient-storing organ. Hypocotyl internode short to long (sometimes modified into nutrient-storing organ), or absent. Mesocotyl present or absent. Coleoptile present (sometimes modified into plumule envelope), with or without lamina, or absent. Collar rhizoids or collar roots sometimes present. Radicula unbranched, usually poorly developed, contractile, persistent or ephemeral (rarely absent). First leaf usually orientated at 180o to plane of cotyledon (in most angiosperms orientated at 90o to plane of cotyledons).

Cytology Membrane complexes absent. Paracrystalline bodies with closely spaced subunits.

DNA Duplication producing monocotyledonous nuclear genes LOFSEP and FUL3 (latter duplication of gene AP1/FUL). Nuclear genes PHYA, PHYB and PHYC present. Nuclear gene PHYE lost. AP3 expression localised on tepal edges. Mitochondrial genes rpl2 and sdh3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, isorhamnetin, syringetin, etc.), O-methylated flavonols, laricitrin, flavonol glycosides, dihydroflavones, flavones, flavone glycosides, flavonoid sulfates, biflavonoids, isoflavones, biflavones, flavanones, homoisoflavanones, hydroxyflavonoids, aurones, luteolin, apigenin, cyanidin, delphinidin, pelargonidin, malvidin, etc., anthoxanthins, deoxyanthocyanins, ethereal oils consisting of mono-, di-, tri- and sesquiterpenes, proanthocyanidins, phenylpropanoids and related curcumins (diarylheptanoids), caffeic acid, chalcones, 6-hydroxyapigenin methyl ethers, diterpenes, triterpenes, oxidized tetracyclic and pentacyclic diterpenes and triterpenes, sesquiterpenes, chalcones, (ent-)epicatechin-4 (non-hydrolyzable tannin), protocatechinic aldehyde, catechins, proanthocyanidins, cinnamic acid, daphnetin, juncosol, caffeic acid derivatives (including caffeic acid sulfate), chlorogenic acid, phenolic glycosides, polyphenolic glycosides with caffeic acid, phenolic sulfates (sulfonated phenolic acids), phenols, norbelladine alkaloids (toxic tyrosine derivatives), isoquinoline alkaloids (benzylisoquinoline alkaloids rare), tryptophane- or tyramine-derived alkaloids, pyrrolizidine alkaloids as 1-aminopyrrolizidine derivatives, indole alkaloids, steroidal alkaloids, quinines, polyhydroxyalkaloids, piperidine alkaloids, lactone alkaloids (tuberostemonine), tropane alkaloids, cholestane glycosides (cardiotoxic bufodienolides, cardenolide glycosides and spirostanol glycosides), tyrosine-derived cyanogenic glycosides etc., steroidal saponins and sapogenins, chelidonic acid, cyanogenic compounds (e.g. cyanogenic glycosides), chrysazine, anthrones (e.g. anthrone-C-glycoside in leaves), tetrahydroanthracenones, naphthoquinones, quinonoid pigments, resins, aromatic acids and ketones, benzoic quinones, shikimic acid- or polyacetate-derived arthroquinones, nepodin, dianellidin, stypandrol, dianellidone, magniferin (glycosylic xanthone), chromones, lactone, phenylpyrones, phenylphenalenones (perinaphthenones), arylphenalenones, phenolic amines, polyamines, acetidine carbonic acid, ascorbic acid, tuliposides (glucose esters), allyl sulfides, allyl disulfides, propyl sulfides, vinyl disulfides, alliin, propionaldehyde, propionthiol, hydroxycinnamic acid, eicosanyl arachidate, crocein, phytosterols, 5-alkylic- and 5-alkenylic resorcinols, homogentisic acid and their glycosides, p-coumaric acid, -sitosterol, ceryl alcohol, amines, non-protein amino acids (tricine [zwitterionic amino acid], S-methylcysteine, etc.), meta-carboxysubstituted aromatic amino- and γ-glutamic peptides, acaroid resins, oxypipe colanic acid (polysaccharide), saccharose esters of diferulic or triferulic acid, and stem fructans present. p-hydroxybenzaldehyde (lignin component) at least often present. Ellagic acid, ellagitannins, lignans, and neolignans not found. Triterpene saponines rare. Hemicelluloses present as xylans.

Fossils Monocotyledon fossils are often difficult to distinguish from other basal angiosperm groups. However, numerous fossils more or less similar to extant Liliidae have been described during the last decades. Many of these have not been assigned to any particular extant clade.

- The oldest known fossil Liliidae are 120–110 My old and resemble Pothooideae (Araceae).

- Acaciaephyllum from the Potomac Group of eastern North America represents herbaceous plants with sheathing leaf bases and an acrodromous reticulate venation. The taxonomic affiliation is highly questioned, although they have been assigned to the monocotyledon stem group (Doyle 1973, etc.).

- A number of fossilized leaves and stems of monocotyledons have been found in Turonian layers in Israel and in the Maastrichtian Deccan Intertrappen Beds of India. These include Geonomites, Limnobiophyllum dentatum, Plumafolium bipartitum, Pontederites eichhornioides, Potamogetophyllum mite, Quturea fimbriata, Typhacites negevensis, Aerophyllites intertrappea, and Aerorhizos harrissii.

- Spinizonocolpites is a Late Cretaceous pollen fossil strongly resembling the extant Nypa (Arecaceae). Early Cretaceous pollen types which have been referred to monocotyledons due to their characteristic monocot morphology include the sometimes frequently occurring Liliacidites and Similipollis.

- Shuklanthus superbum is a racemose inflorescence found in the Maastrichtian layers of the Indian Deccan Intertrappean Beds, whereas Viracarpon from the same layers represents an infructescence that may actually belong to the same species. The unisexual flowers are trimerous with six tepals and six uniovulate carpels. The fruits consist of single-seeded drupelets.

- Deccananthus savitrii from the Deccan Intertrappean Beds of India is a trimerous flower with six tepals and six stamens. The pollen grains are trichotomosulcate and the ovary is trilocular.

- Tricoccites trigonum comprises three-seeded trilocular drupes, likewise from the Deccan Intertrappean Beds, resembles extant Arecaceae and Pandanaceae.

- Eriospermocormus indicus is a fossilized corm from the Deccan Intertrappean Beds. It is somewhat similar to the Eriospermum (Ruscaceae), but its systematic affiliation is uncertain.

Systematics Acorus is sister to the remaining monocots and Alismatales successive sister-group to the remainder.

A widely accepted hypothesis is that Liliidae have evolved from helophytic (or even aquatic) ancestors. Numerous adventitious roots replacing an ephemeral main root (instead of a single tap-root), sympodial growth, atactostele (vascular bundles scattered in stem), absence from normal secondary lateral growth, and usually linear leaves lacking normal lamina are characteristics of the monocots which have been explained through this hypothesis. Moreover, most members of the two basal monocot clades, Acorus and Alismatales, are helophytes or aquatic.

Reticulately veined pseudolamina and baccate fruits – probable adaptations to forest habitats – have evolved in parallel in many monocotyledon clades. According to Givnish & al. (2005), baccate or drupaceous fruits have evolved 21 times and reticulodromous venation perhaps between 25 and 30 times during the evolution of Liliidae.

Phylogeny of Liliidae based on DNA sequence data (Tamura & al. 2004; Chase & al. 2006; Graham & al. 2006; Soltis & al. 2011). Acorus and Alismatales are supported by 100% (bootstrap-value) as successive sister-groups to the remaining Liliidae. The clade [Pandanales+Taccales] also has very high support. Liliales and Iridales are successive sisters to the remainder, the Commelinidae (a clade bootstrap support of 100%)

Nartheciidae S. W. Graham et W. S. Judd in Taxon 56: E25. Aug 2007

[Alismatales+[Petrosaviaceae+[Taccales+Pandanales]+[Liliales+[Iridales+Commelinidae]]]]

Potential synapomorphies: Ethereal oils absent. Raphides present. Ptyxis (of pseudolamina) supervolute-curved or variations of this principle. Endothecium developing directly from undivided outer secondary parietal cells. Pollen grains boat-shaped. Sexine reticulate. Tectum with finer sculpture at pollen ends. Endexine absent. Septal (epithelial) nectaries often present (intercarpellary fusion often postgenital). – Nartheciidae contain all monocotyledons except Acorus.

Petrosaviidae S. W. Graham et W. S. Judd in Taxon 56: E25. Aug 2007

[Petrosaviaceae+[Taccales+Pandanales]+[Liliales+[Iridales+Commelinidae]]]

Potential synapomorphies: Pseudolamina developing from leaf base zone. Epidermis with bulliform cells. Stomata anomocytic. Cuticular wax crystalloids sometimes as parallel platelets. Colleters (squamulae intravaginales) absent. Starch grains simple, amylophobic. Cyanogenic glycosides infrequent. – Petrosaviidae comprise Liliidae except Acorus and Alismatales.

[[Taccales+Pandanales]+[Liliales+[Iridales+Commelinidae]]]

Potential synapomorphies: Nucellar cap absent. Endosperm development nuclear.

Pandananae Thorne ex Reveal in Novon 2: 236. 1992

[Taccales+Pandanales]

Potential synapomorphy: Outer integument two (or three) cell layers thick.

[Liliales+[Iridales+Commelinidae]]

Synapomorphies other than from DNA sequences have not been found.

[Iridales+Commelinidae]

Potential synapomorphy: Style long.

Commelinidae Takht., Sist. Filog. Cvetk. Rast.: 514. 4 Feb 1967 [sensu S. W. Graham et W. S. Judd]

[Arecaceae+Dasypogonaceae+[Cyperales+Commelinanae]]

Potential synapomorphies: Unlignified cell walls containing UV-fluorescent ferulic acid and coumaric acid (fluorescence of unlignified cell walls caused by presence of ferulic and/or coumaric acids). Vessel elements sometimes present in stem and leaves. Cells with silica bodies present in leaves. Stomata paracytic or tetracytic. Cuticular wax crystalloids sometimes Strelitzia type (as aggregated rodlets resembling scallops of butter). Inflorescence indeterminate. Peduncle bracteate. Filaments adnate to inner tepals (epipetalous). Pollen grains containing starch. Embryo short and wide. Expression of B-class gene orthologue of nuclear gene PISTILLATA probably restricted to androecium and inner tepals.

[Cyperales+[Commelinales+Cannales]]

Potential synapomorphies: Primary cell walls usually with glucurono-arabinoxylans. Stomatal subsidiary cells with parallel divisions. Endosperm with starch.

[Commelinales+Cannales]

Potential synapomorphies: Inflorescence a helicoid cyme (with many-flowered cincinnal branches). Tapetum amoeboid-periplasmodial or invasive. Operculum of seed combined with micropylar collar (micropylar collar formed at apex of outer integument, protruding as orbicular wedge into megasporangium; sclerotic layer of seed coat discontinuous at site of micropylar collar; collar sometimes with special rupturing layer). Endotestal cells silicified.

[Ceratophyllum+Tricolpatae]

Potential synapomorphy: Ethereal oils absent. – Ceratophyllum is sister to Tricolpatae in the maximum-likelihood tree of Soltis & al. (2011), but recovered as sister-group to Liliidae in the maximum-parsimony tree of the same study.

Ceratophyllum has a large number of autapomorphies, in part due to their highly specialized aquatic lifestyle. Roots, vessels, stomata and cuticular waxes are absent and the perianth is reduced. Even their pollen morphology – inaperturate (to indistinctly monocolpate) and with very reduced exine – may be a result of adaptation to an aquatic environment. The microsporogenesis is successive (like in monocots) in some species and simultaneous (like in most eudicots) in others. Further investigations of the pollen development in Ceratophyllum are certainly critical to our understanding of pollen character optimization among Tricolpatae.

Ceratophyllum share many features with the majority of Liliidae, including ephemeral primary root, closed stem vascular bundles, absence of interfascicular cambium, absence of vessels in stem and leaves, perianth (if present) trimerous, and successive microsporogenesis. On the other hand, it seems to be very difficult to find morphological synapomorphies for the clade [Ceratophyllum+Tricolpatae].

Tricolpatae M. J. Donoghue, J. A. Doyle et P. D. Cantino in Taxon 56: E26. Aug 2007 (tricolpates, eudicots)

Potential synapomorphies: Root epidermis derived from root cap. Nodes 3:3 (trilacunar with three leaf traces). Foliar lamina usually developing from leaf apex. Stomata anomocytic. Cuticular wax crystalloids as clustered tubuli (Berberis type), with nonacosan-10-ol as dominating wax. Chloranthoid leaf teeth possibly apomorphous (also in Chloranthaceae). Flowers cyclic, sometimes dimerous. Outer tepals (sepals) with three traces. Inner tepals (petals) with one trace. Stamens few, individually antetepalous (also in Lauraceae). Polyandry (secondary) widespread. Initial primordia sometimes five, ten or annular, sometimes centrifugally developing. Filaments fairly slender. Anthers basifixed. Microsporogenesis simultaneous. Pollen tetrads tetrahedral. Pollen grains triaperturate. Apertures in pairs at six points on young tetrad, according to Fischer’s rule. Cell division (cleavage) centripetal. Pollen wall with endexine. Carpels with complete postgenital fusion. Style solid (not hollow). x = 7. Vacuolar crystal formation associated with membranes and paracrystalline bodies with widely spaced subunits. Myricetin and delphinidin scattered, asarone absent (present in some asterids).

Fossils The oldest known fossil tricolpate pollen grains have been found in Late Barremian to Early Aptian strata in England, Portugal, Israel, Egypt, tropical West Africa, and eastern North America. The exine is finely to coarsely reticulate or striate, with a columellate infratectum. Examples of early fossils not assigned to any particular eudicot clade are as follows.

- Sinocarpus decussatus comprises parts of infructescences and leaves from the Aptian of China. The decussate leaves are provided with chloranthoid teeth and the fruits are formed by three or four whorled and partially connate carpels.

- Hyrcantha karatscheensis is represented by reproductive axes and fruits from the mid-Albian of Kazakhstan. The gynoecium is composed of three to five free carpels.

- Ternariocarpites floribundus is an infructescence with free carpels from the Albian of the Russian Far East. The fruitlets are follicular and the five tepals are persistent.

- Ranunculaecarpus quinquecarpellatus from the Albian of East Siberia consists of an apocarpous fruit with five follicular carpels. The fossilized bicarpellate syncarpous fruit of Araliaecarpum kolymense emanates from the same Siberian locality.

- The Cenomanian flower Callicrypta chlamydea from eastern Siberia has a perianth consisting of three whorls, stamens/staminodia and six free carpels.

- Cathiaria zhilinii is known from several localities from eastern Europe to Japan. It comprises fruiting structures with monocarpellate single-seeded fruits from the Cenomanian to the Coniacian.

- Numerous follicular fruits (Agapitocarpus emisxus, Chontrocarpus pachytoichus, Maiandrocarpus moirasmenus, Malliocarpus batrachoides, Mitocarpus elegans, Xylocarpus rhitidodes, Zeugarocarpus) have been found in Late Santonian to Early Campanian layers of Sweden. The different fossil species are relatively similar to each other. The gynoecium is apocarpous or monocarpellate, the carpels are plicate and the multiple ovules, when known, are anatropous and bitegmic. Traces of tepals are absent.

Systematics Apart from the unresolved relationship (beyond Ranunculaceae) between Proteales, Sabiaceae and the remaining Tricolpatae, Sabiaceae have been recovered as sister to Proteales in several studies (Qiu & al. 2006; Moore & al. 2008; Burleigh & al. 2009; Moore & al. 2011; Soltis & al. 2011), yet with weak to moderate support. Hence, I am apt to include Sabiaceae in Proteales. In other analyses, Sabiaceae have been identified (with weak support) as sister to Tricolpatae except Ranunculales and Proteales (Soltis & al. 2008), to Tricolpatae except Ranunculales (Worberg & al. 2007; Qiu & al. 2010; etc.), or even as sister to Buxales (Kim & al. 2004). The recovered sister-group relationships in this part of Tricolpatae largely depend on number and types of sequenced genes (plastid and/or mitochondrial and/or nuclear genes).

In all, I have followed Moore & al. 2010, Moore & al. 2011 and Soltis & al. 2011, since there phylogenies are strongly supported.

[Proteales+Sabiaceae+[Trochodendrales+[Didymelales+Gunneridae]]]

Potential synapomorphy: Axial/receptacular nectaries sometimes present.

[Trochodendrales+[Didymelales+Gunneridae]]

Potential synapomorphies: Mitochondrial gene rps2 absent (lost). Benzylisoquinoline alkaloids absent.

Gunneridae D. E. Soltis, P. S. Soltis et W. S Judd in Taxon 56: E27. Aug 2007

[Gunnerales+Pentapetalae]

Potential synapomorphies: Leaf margin serrate. Compitum present. Duplication of floral organ identity B-class gene paleoAP3 (yielding euAP3 and TM6 paralogs). PI-dB motif present. Small deletion in the 18SrDNA frequently present. Ellagic and gallic acids abundant.

Fossils In the Late Albian to the Early Cenomanian of Nebraska, there are unambiguous fossils of pentamerous heterochlamydeous flowers (with differentiated calyx and corolla) which are assignable to the Gunneridae.

Systematics Several other gene duplications seem to have taken place in the ancestors of either Gunneridae or Pentapetalae (or sometimes even earlier), i.a. duplication of nuclear floral regulatory genes AP1/FUL or FUL-like gene (yielding euAP1, euFUL and AGL79); duplication of nuclear gene RPB2; duplication of AG-like C-class gene (yielding PLE and euAG paralogs); duplication of nuclear genes AGL2/3/4 (yielding SEP1 and FBP6) and AGL1/2/3, etc. (see, i.a., Kramer & Zimmer 2006; Kramer & al. 2004; Kramer & al. 2006). The knowledge of many of these duplications is insufficient for many critical clades (Proteales, Sabiaceae, Trochodendrales, Didymelales, Gunnerales, Dilleniaceae, Santalales, Berberidopsidales, etc.). Consequently, it is still impossible to optimize them convincingly on the tree.

Since Gunnerales are sister to Pentapetalae, detailed knowledge of floral development in Gunnera and Myrothamnus is important for our interpretation of floral characters in the crown group of Tricolpatae. The flowers of Gunnerales are strongly adapted to wind pollination, a fact that makes it even more difficult to draw conclusions on homologies. Furthermore, the organization of the androecial and perianth whorls are more similar to basal Tricolpatae than to Pentapetalae.

Pentapetalae D. E. Soltis, P. S. Soltis & W. S. Judd in Taxon 56: E27. Aug 2007

Potential synapomorphies: Root apical meristem closed. Flowers pentamerous, with whorled floral parts. Calyx/sepals and corolla/petals distinct. Sepals enclosing flower in bud (sepals and petals encircling floral axis). Sepals with three or more traces. Petals with one trace. Nectariferous disc present. Stamens twice the number of sepals/petals (sometimes numerous, but then usually fasciculate), developing internally/adaxially to corolla whorl and successively alternating from five (ten) primordials, and/or centrifugally. Pollen grains tricolporate. Carpels five (although three also frequent; when carpels two, then superposed). Style present. Stigma not decurrent. Placentation axile. Endosperm development nuclear. Fruit dry, dehiscent (when capsule then loculicidal). RNase-based gametophytic incompatibility system present (stylar response mediated by glycoprotein with RNase activity). Whole genome triplication (γ triplication) leading to paleohexaploidy. Cyanogenesis also via phenylalanine, isoleucine or valine pathways (cyanogenic compounds also phenylalanine-, isoleucine- or valine-derived). – Numerous stamens have evolved multiple times in Pentapetalae. In these cases the stamens are often arranged in fascicles. They may develop – centripetally or centrifugally – from usually five antepetalous or ten separate primordial or from an annular androecial primordium.

[Dilleniaceae+[Santalales+[[Caryophyllales+Berberidopsidales]+Asteridae]]]

Synapomorphies other than from DNA sequences have not been found for this clade. – The position of Dilleniaceae is still very unstable. They were recovered as sister-group to the clade [Saxifragales+Rosidae] (Superrosidae) in an analysis of complete plastid genome sequence data (Moore & al. 2010). Presence of stipules (usually inserted on the stem/branch) is a feature common to Dilleniaceae and rosids. On the other hand, in an analysis of inverted repeat sequences, Dilleniaceae were identified as sister to the clade [Superasteridae+Superrosidae], i.e. to all other Pentapetalae (Moore & al. 2011). In the analyses by Soltis & al. (2011), Dilleniaceae were sister-group to Superasteridae in the maximum-likelihood analysis, whereas the maximum-parsimony analysis revealed them as sister to Caryophyllales. A position as sister to Caryophyllales was revealed by Bell & al. (2010) and to Superasteridae by Arakaki & al. (2011). Qiu & al. (2010) recovered the topology [[Dilleniaceae+ Berberidopsidales]+[Asteridae+[Caryophyllales+Santalales]]]. Pending more convincing information I leave Dilleniaceae as sister to Superasteridae, even if the support for the position is not very strong.

Superasteridae W. S. Judd, D. E. Soltis et P. S. Soltis in Amer. J. Bot. 98: E23. Apr 2011

[Santalales+[[Caryophyllales+Berberidopsidales]+Asteridae]]

Potential synapomorphies: Corolla in Santalales and Asteridae visible in bud stage long before floral dehiscence (possibly a parallelism).

[Caryophyllales+Asteridae]

Potential synapomorphies: Seed coat exotestal. Embryo elongated. – Presence and morphology of transfer cells and intermediary cells may provide synapomorphies.

Asteridae Takht., Sist. Filog. Cvetk. Rast.: 405. 4 Feb 1967

[Loasales+[Ericales+Gentianidae]]

Potential synapomorphies: Tension wood not frequent. Petals connate (sometimes only visible during early development). Corolla enclosing androecium and gynoecium in bud. Nectary gynoecial, supplied from gynoecial traces. Style single, long. Ovules unitegmic. Integument thick. Endothelium present. Megasporangial epidermis not persistent. Exotestal cells lignified, especially on anticlinal and/or inner periclinal walls. Endosperm development cellular. Embryo elongated. Nicotinic acid metabolized to its arabinosides. Iridoids and/or polyacetylenes frequent in many (also basal) clades. – Early initiation of the corolla tube may be a synapomorphy at this level.

Vegetative anatomy Ectomycorrhiza sometimes present. Medullary and/or cortical vascular bundles (sometimes inverted) often present. Secondary lateral growth normal, anomalous (via concentric cambia or cylindrical cambium) or absent. Vessel elements usually with scalariform or simple (rarely reticulate) perforation plates; lateral pits alternate, opposite or scalariform, simple or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements fibre tracheids, libriform fibres or tracheids with simple or bordered pits. Wood elements sometimes storied. Tyloses sometimes frequent. Palisade mesophyll sometimes with arm cells. Secondary phloem often stratified into hard fibrous and soft parenchymatous layers. Intraxylary phloem rarely present. Sieve tube plastids usually Ss type (sometimes Pc, Pcs or Pcf type). Non-articulated branched or unbranched laticifers with white or bluish latex sometimes present. Schizogenous secretory canals and cavities with ethereal oils, resins, gutta-percha or mucilage sometimes present. Heartwood sometimes with gum-like substances. Sclerenchymatous idioblasts often present. Silica bodies sometimes present in parenchyma cells. Calciumoxalate as raphides, styloids, druses, crystal sand, or acicular, rhomboidal or prismatic crystals often present (rarely sphaerites or tuberculate, glochidiate or stinging hairs).

Trichomes Hairs unicellular or multicellular, simple or branched, furcate, stellate, fasciculate, dendritic, candelabra-, funnel- or cup-shaped, T-shaped, arachnoid, peltate, lepidote, vesicular, flagellar hairs, bristle- or prickle-like, or papillose (rarely moniliform), or absent; stalked or sessile glandular hairs often present; laticiferous hairs sometimes present.

Leaves Alternate or opposite (rarely verticillate), pinnately or palmately compound or simple and entire or lobed, with conduplicate, involute, supervolute, convolute, revolute, curved or flat ptyxis. Stipules usually absent (sometimes cauline or interpetiolar, rarely intrapetiolar); leaf sheath usually absent (sometimes present, conspicuous). Colleters often present. Venation pinnate or palmate, eucamptodromous, brochidodromous, craspedodromous, or semicraspedodromous (rarely parallelodromous, flabellate, actinodromous or acrodromous), or leaves single-veined. Stomata usually anomocytic, paracytic, or diacytic (sometimes diallelocytic, anisocytic, helicocytic, tetracytic, laterocytic or cyclocytic). Cuticular wax crystalloids as platelets, scales or rodlets (rarely threads or amorphous or as tubuli dominated by diketones). Domatia as pits, pockets or hair tufts, or absent. Secretory canals and cavities sometimes with ethereal oils, resins, gums, or mucilage. Epidermis with or without mucilage cells. Secretory oil cavities occasionally present. Mesophyll often with resiniferous, laticiferous or sclerenchymatous idioblasts. Mesophyll cells often with calciumoxalate as prismatic crystals or druses (sometimes sphaerites, styloids, raphides, acicular crystals or crystal sand). Leaf margin entire, serrate, crenate or lobate. Leaf teeth often with glandular apex, one accessory vein proceeding into tooth, the other running above tooth; or with simple vein and translucent caducous apex; leaf teeth sometimes theoid.

Inflorescence Panicle, fascicle, raceme-like, corymb, cincinnate, scorpioid, spicate, catkin-like, thyrsoid, botryoid, umbellate or capitate thyrse, often as whorls of dichasial and/or monochasial partial inflorescences, or racemes, spikes, umbels or heads (flowers sometimes solitary or paired, axillary). Floral prophylls (bracteoles) usually pairwise, lateral (rarely absent).

Flowers Actinomorphic or zygomorphic (rarely resupinate or asymmetrical). Pedicel sometimes articulated. Hypogyny, epigyny or half epigyny. Sepals (two to) four or five (to 16), with imbricate, valvate, contorted or open aestivation, usually more or less connate. Petals (three or) four or five (to 18), with imbricate, valvate, contorted, convolute or ascending-cochlear (rarely induplicate, descending-cochlear or open) aestivation, free from each other or connate into campanulate, hypocrateromorphous, discoid, urceolate, tubular or infundibuliform, often bilabiate corolla, sometimes spurred. Nectaries usually present on petal bases or intrastaminal nectariferous disc, usually annular (sometimes cupular, rarely unilateral), entire or lobate, or as separate nectariferous glands alternating with stamens, often only on abaxial side (rarely absent).

Androecium Stamens (two to) four or five (to 16, rarely to more than 1.200), usually in one whorl, haplostemonous, antesepalous, alternipetalous (rarely in two or several whorls). Filaments usually free from each other (sometimes more or less connate), usually adnate to petals/corolla tube (epipetalous). Anthers usually free from each other (sometimes more or less connate, occasionally adnate to style forming gynostegium), basifixed, ventrifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (sometimes disporangiate or synthecal, rarely monosporangiate or octosporangiate), usually extrorse or introrse (rarely latrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical or basal pores). Placentoid often present. Tapetum secretory or amoeboid-periplasmodial. Staminodia one to three or absent; female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains 2–3(–10)-colpate, -porate, or -colpor(oid)ate (sometimes syncolpate, sometimes with few or numerous pseudocolpi; rarely monocolpate, pororate, pantoporate, inaperturate, pericolpate, stephanocolp[or]ate or spiraperturate), usually shed as monads (rarely dyads, triads, tetrads or polyads), bicellular or tricellular at dispersal. Exine with usually columellate (sometimes granular) infratectum.

Gynoecium Pistil composed of usually two (rarely up to c. 30) connate carpels (rarely one carpel). Ovary superior, inferior or semi-inferior, unilocular or bilocular (sometimes incompletely septate, sometimes pseudomonomerous, rarely up to c. 30-locular), sometimes with locules divided by secondary septa. Style usually single, usually terminal (sometimes gynobasic), simple or lobate (stylodia sometimes two, free), or absent. Stigma capitate, truncate, discoid, infundibuliform, peltate, clavate, fusiform, or lobate, or stigmas punctate to capitate, papillate or non-papillate, usually Dry or Wet type. Pistillodium usually absent (male flowers often with pistillodium).

Ovules Placentation apical, subapical, basal, subbasal or axile (sometimes free central or parietal, rarely intrusive). Ovules one or two to numerous (to several hundred) per carpel, anatropous, hemianatropous, amphitropous, pleurotropous or campylotropous (rarely orthotropous, circinotropous or hypertropous), ascending, horizontal or pendulous, apotropous or epitropous, usually unitegmic (sometimes bitegmic, rarely ategmic), usually tenuinucellar (sometimes crassinucellar), often non-vascularized. Integument when single often with dermal origin; outer integument when present usually subdermal in origin. Placental obturator often present. Hypostase often present. Endothelium usually present. Archespore sometimes multicellular. Parietal cell often dividing. Nucellar cap present or absent. Megagametophyte usually monosporic, Polygonum type (rarely Oenothera type, or disporic, Allium type, or tetrasporic, Adoxa, Penaea, Fritillaria, or Drusa type). Synergids sometimes with filiform apparatus. Antipodal cells sometimes persistent, sometimes proliferating. Endosperm development usually cellular (sometimes nuclear, rarely seemingly helobial or other aberrant type). Endosperm haustorium chalazal and/or micropylar, or absent. Embryogenesis onagrad, asterad, solanad, chenopodiad or caryophyllad.

Fruit A loculicidal and/or septicidal capsule (sometimes septifragal, rarely a pyxidium, a denticidal capsule or irregularly dehiscing), a drupe, a berry, an achene, or a schizocarp with usually nutlike (rarely baccate, drupaceous or follicular) mericarps (rarely a samara or syncarp).

Seeds Perisperm not developed. Endosperm copious to sparse or absent, with starch, oil, hemicellulose and/or proteins (sometimes with petroselinic acid). Embryo straight to curved, sometimes oily, well to poorly differentiated, usually without chlorophyll. Cotyledons usually two (rarely one, four or absent), accumbent or incumbent, rarely foliaceous.

Cytology Protein bodies often present in mesophyll cell nuclei.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, gossypetin), flavones, methyl flavones, sulfated or O-methylated flavonoids, oxygenated flavones, flavone glycosides, catechins, cyanidin, delphinidin, acylated anthocyanins, davidigenin, Route I carbocyclic iridoids, Route II iridoids (also C4-decarboxylated iridoids, i.e. iridoid glycosides and glycosides of monoterpenoid lactones), Group I carbocyclic iridoids, Group II carbocyclic iridoids, Group III carbocyclic iridoids, Group IV carbocyclic iridoids, Group V carbocyclic iridoids, Group VI secoiridoids, Group VII secoiridoids, Group VIII secoiridoids, Group IX secoiridoids, Group X secoiridoids (including iridoid pyridine alkaloids), iridoid aldehydes, iridoid glycosides (usually C10 and C11 iridoid glycosides, rarely C8 iridoid glycosides), iridoid coumarins, cardenolides, sarracenin, oleanolic acid derivatives, monoterpenes, diterpenes, sesquiterpenes, triterpenes, ethereal oils, balsams, and resins consisting of monoterpenoids, triterpenoids, sesquiterpenoids, phenylpropanoids, etc., free terpenic acids, dammaranes, ursolic acid, arjunolic acid derivatives, ellagic and gallic acids, ellagitannins, non-hydrolyzable tannins, proanthocyanidins, ursolic and caffeic acid, ursolic acid and caffeic acid esters (chlorogenic acid often abundant), nicotinic acid compounds, quinazolinone alkaloids, benzylisoquinoline and other isoquinoline alkaloids, hemlock alkaloids, C17 and other indole alkaloids, pyrrolizidine and pyrrolidine alkaloids, pyrrolizidine alkaloids as macrocyclic diesters or as aliphatic monocarboxylic esters or esters of arylic or aralkylic acids, strongly toxic diterpene alkaloids, steroid alkaloids, tryptophane-derived alkaloids, hygroline alkaloids, nortropane- and tropane-3α-ols, tropane-3β-ols, tropane alkaloids, hydroxyl tropines, toxic (cardiotonic) steroid glycosides, syringin, phenolic heterosides, 3-galactoside, triterpene saponins, simple coumarins (e.g. umbelliferone), hydroxycoumarins, pyranocoumarins, dihydropyranocoumarins, furanocoumarins, dihydrofuranocoumarins etc., p-coumaride, betaines, hydroxycinnamic acids (ferulic acids), coniferin, silicic acid, polyacetate-derived arthroquinones, shikimic acid derived arthroquinones, benzoquinones and naphthoquinones (also prenylated) and their derivatives, anthraquinones, terpenoid-derived quinones, phenylalanine- or tyrosine-derived cyanogenic compounds, cyanogenic compounds (i.a. cyanogenic glycosides), simmondsinoid compounds, arbutin, eleostearic acid, valerianic acid, actinidin, acetophenones, sesquiterpene lactones, withanolide steroidal lactones, phenantrenes, phenylpropenes (e.g. myristicin), germacrane-like compounds, asarone, syringaresinol, pinoresinol, aliphatic monocarboxylic acids, polyacetylenes (e.g. falcarinone) derived from fatty acids, mostly aliphatic C17 acetylenes, petroselinic (cis-6-octadecenoic) acid (in endosperm), xanthones, eucommin A, benzopyrones, cucurbitacins, dihydrosterculic acid, lignans, myo-inositol, and gutta-percha. Carbohydrates often stored as oligosaccharides (i.a. stachyose, mannitol, hexites, saccharose, or umbelliferose). Carbohydrates sometimes stored as oligo- or polyfructosans (inulin) with kestose or isokestose linkages (starch then usually absent).

Systematics The combination of solitary and very long vessel elements (800 µm or more) with scalariform perforation plates, usually opposite pitting of vessel elements, non-septate and very long fibres (2.190 µm or more) with bordered pits, and axial parenchyma diffuse or diffuse-in-aggegates and paratracheal scanty (the Baileyan wood anatomical syndrome) is abundant among asterids. Compound leaves are fairly uncommon and the leaflets are often articulated and/or distinct. Stipules are also relatively infrequent.

Sympetalous zygomorphic flowers are often combined with epipetalous stamens. Sympetaly is a character common to the majority of Asteridae. Even many clades with choripetalous flowers (possessing free petals) seem to be principally sympetalous, since they develop an early annular primordium and show early initiation of the corolla tube (Leins & Erbar 2003; Erbar & Leins 2011). Early initiation of the corolla tube occurs particularly among Campanulidae (i.a. in Araliales, Campanulales and Dipsacales), but also in some Lamiidae (e.g. Oleaceae among Plantaginales, and Rubiaceae in Rubiales) and in several Loasales. Unfortunately, knowledge about corolla initiation in the basal clades of Gentianidae is lacking.

The ovules in Asteridae are usually characterized as tenuinucellar and unitegmic, although the seemingly single integument may in fact be composed of two fused integuments. In bitegmic asterids the outer integument usually has a subdermal origin. The integument in unitegmic asteroids are often dermal in origin and possibly corresponds to the inner integument in most bitegmic angiosperms.

Phylogeny of Asteridae based on DNA sequence data (González & al. 2007; Winkworth & al. 2008; Tank & Donoghue 2010; Soltis & al. 2011; etc.). Cornales and Ericales are successive sister-group to the remainder with high bootstrap support (97% and 100%, respectively, in Soltis & al. 2011). The two major clades are Lamiidae (Icacinaceae to Solanales) and Campanulidae (Aquifoliales to Dipsacales). Aquifoliales are sister to the remaining Campanulidae – Apiidae – with high support, yet the sister-group relationships within Apiidae are unclear. Bruniaceae are sometimes sister to Columelliaceae, with a bootstrap support of 79% in plastid DNA analyses, and <50% in combined analyses (Soltis & al. 2011). A likewise weakly supported Dipsapiidae clade (Tank & Donoghue 2010) is sometimes recovered (Winkworth & al. 2008; Soltis & al. 2011; etc.), consisting of Araliales as sister to Paracryphiaceae and Dipsacales. Escalloniaceae are recovered as sister to Campanulales, with low support, in some studies (e.g. Soltis & al. 2011), yet sister to all Campanulidae except Aquifoliales in, i.a., Tank & Donoghue (2010).

[Ericales+Gentianidae]

Potential synapomorphies: Ovules without parietal tissue, tenuinucellar.

Gentianidae R. G. Olmstead, W. S. Judd et P. D. Cantino in Taxon 56: E29. Aug 2007

Potential synapomorphies: Sugar transport in phloem active. Inflorescence basically cymose. Petals connate, forming distinct tube. Stamens as many as (or less than) sepals/petals, antesepalous. Polyandry when present usually associated with increased numbers of petals or carpels. Pollen grains sometimes with orbicules (Ubisch bodies). Filaments adnate to petals (epipetalous). Duplication of nuclear gene PI. Non-hydrolyzable tannins infrequent. Ellagic acid absent.

Ellagic acid occurs together with iridoids in many basal asterid clades, especially in Loasales and Ericales, whereas ellagic acid is absent in Gentianidae. Moreover, polyandry (possession of numerous stamens), a characteristic feature of several clades in Loasales and Ericales, is very infrequent among Gentianidae (rare examples are Hoplestigma in Boraginaceae and Dialypetalanthus in Rubiaceae). Polyandry may be correlated with increase in tepal (and sometimes carpel) number, contrary to the case in non-asterid Tricolpatae.

Garryidae R. G Olmstead, W. S. Judd et P. D. Cantino in Taxon 56: E30. Aug 2007

Synapomorphies other than from DNA have not been found. – Garryales, Icacinaceae, Emmotaceae, the Apodytes clade and Cassinopsis form a basal grade in Garryidae. Oncotheca and Metteniusa, formerly sometimes believed to be closely allied to Garryales or Icacinaceae, form a basal grade (or perhaps clade) in Lamiidae. – Garrya and Eucommia have only the d copy of the gene RPB2, whereas RPB2 is duplicated in Ilex.

Lamiidae Takht. ex Reveal in Phytologia 74: 178. 25 Mar 1993 [sensu R. G. Olmstead et W. S. Judd] (euasterids I)

Potential synapomorphies: Carpels two, superposed. Introns 18–23 in d copy of gene RPB2 lost. Caffeic acid present. – The I copy of the gene RPB2 is present in most Lamiidae and also in Ericales (absent from Rosidae and remaining Asteridae).

[Oncothecaceae+Metteniusaceae]

Potential synapomorphies: Vessel elements with scalariform perforation plates. Nodes 5:5. Petiole vascular bundle transection arcuate, complex. Bracts thick, triangular. Stamens as many as petals, alternipetalous. Anthers basifixed. Carpels five. Ovules two per carpel. Funicle long. Fruit a drupe, with persistent calyx. Endosperm copious.

[Boraginaceae+Vahliaceae+[Rubiales+[Plantaginales+Solanales]]]

Potential synapomorphies: Vessel elements with simple perforation plates. Corolla tube initiation late (petals appearing as distinct primordia that subsequently fuse). Nectary (vascularized) at base of ovary. Carpels two. Style long. 8-ring desoxyflavonols sometimes present.

Scandianthus costatus and S. major, from the Late Santonian to the Early Campanian of southern Sweden, may be attributed to some group of Lamiidae. They comprise bisexual actinomorphic epigynous flowers with pentamerous perianth and androecium and a gynoecium formed by two connate carpels. The sepals and petals are free and the stamens diplostemonous. The nectariferous disc is intrastaminal. The pollen grains are tricolporate and tectate, the ovaries unilocular, the styles free, and the capsules many-seeded and apically dehiscent.

[Rubiales+[Plantaginales+Solanales]]

Synapomorphies other than from DNA sequences have not been found.

[Plantaginales+Solanales]

Potential synapomorphies: Nodes 1:1. Sepals connate. Anther theca with placentoid. Endothelium present. Myricetin, iridoids and non-hydrolyzable tannins usually absent. – Protein crystals seem to be frequently present in the nucleus, although the knowledge of their distribution is still very insufficient. Presence of arabino- and galactoxyloglucans (instead of fucogalactoxyloglucans in cell walls may be a synapomorphy.

Campanulidae M. J. Donoghue et P. D. Cantino in Taxon 56: E30. Aug 2007 (euasterids II)

Potential synapomorphies: Vessel elements with scalariform perforation plates. Corolla with valvate aestivation. Corolla tube initiation early; corolla tube starting as annular meristem, ‘ring primordium’, from which separate petals develop (petal primordia originating on ‘ring primordium’ and staminal primordia arising in front of interprimordial connections). Petals often with acuminate apex. Endosperm copious. Embryo short or very short. Myricetin absent. – Information on corolla tube initiation is very insufficient and for numerous clades absent. Epigyny (inferior ovary) may be a synapomorphy here, yet in that case with several reversals to hypogyny. The I copy of the nuclear gene RPB2 is present in at least Escallonia of Escalloniaceae and in Ilex (Aquifoliaceae), but is lost in the majority of Campanulidae. Ilex has also lost the introns 18–23 of the d copy of RPB2, whereas these introns persist in Escallonia. As usual, the situation in the smaller clades (i.a. Bruniales and Paracryphiales) is not known.

Silvianthemum, from the Late Santonian to the Early Campanian of southern Sweden, comprises bisexual actinomorphic epigynous flowers with pentamerous perianth and androecium and a gynoecium of three free carpels. The surface is beset with peltate and simple hairs. The eight or nine stamens are probably inserted in two series. The pollen grains are tricolpate and tectate, the ovaries unilocular, the styles free, and the numerous ovules anatropous and bitegmic. A close relationship with Quintinia in Paracryphiaceae has been suggested.

Apiidae M. J. Donoghue et P. D. Cantino in Taxon 56: E31. Aug 2007

[Campanulales+[Escalloniaceae+[Bruniales+[Araliales+[Paracryphiaceae+Dipsacales]]]]]

Potential synapomorphies: Epigyny. Corolla tube initiation early. Carpels two or three. I copy of gene RPB2 lost. Iridoids or polyacetylenes present. – This clade comprises all Campanulidae except Aquifoliales.

Dipsapiidae D. C. Tank et M. J. Donoghue in Syst. Bot. 35(2): 434. 2010

[Araliales+[Paracryphiaceae+Dipsacales]]

Potential synapomorphy: Nodes 3:3.

Dipsidae D. C. Tank et M. J. Donoghue in Syst. Bot. 35(2): 434. 2010

[Paracryphiales+Dipsacales]

Potential synapomorphies: True tracheids present. Leaf margin serrate. Inflorescence terminal.

Superrosidae W. S. Judd, D. E. Soltis et P. S. Soltis in Amer. J. Bot. 98: E21-E22. Apr 2011

[Saxifragales+Rosidae]

Potential synapomorphies: Stipules present. Nodes 3:3, trilacunar with three leaf traces. Sepals trilacunar and supported by three traces (instead of unilacunar as in many basal Tricolpatae). Nectaries receptacular, supplied from receptacular or androecial traces. I copy of nuclear gene RPB2 lost. – Floral mucilaginous cells with strongly thickened mucilaginous inner periclinal wall and typical cytoplasm may be a synapomorphy of Superrosidae.

Rosidae Takht., Sist. Filog. Cvetk. Rast.: 264. 4 Feb 1967

[Vitaceae+[Fabidae+Malvidae]]

Potential synapomorphies: Mucilage cells with thickened inner mucilaginous periclinal walls and distinct cytoplasm. Anthers articulated (dorsifixed, transition to filament narrow, connective thin). Embryo elongated. Genome duplication leading to paleohexaploidy.

Vegetative anatomy Ectomycorrhiza sometimes present. Root nodules containing nitrogen-fixing endosymbiotic actinobacteria (Frankia) sometimes present. Secondary lateral growth normal, anomalous (from cylindrical cambium or concentric cambia) or absent. Vessel elements with simple or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, scalariform or opposite, simple or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits. Intraxylary phloem sometimes present. Sieve tube plastids Ss, S0, Pc, Pcs or Pcfs type (rarely Pfs type); sieve tubes sometimes with non-dispersive P-protein bodies. Schizogenous secretory cells, canals or cavities or glands with resins, balsam, tannins, ethereal oils or other secretions often abundant. Lysigenic or schizogenic mucilage ducts and cavities sometimes present. Laticiferous cavities or ducts sometimes present. Heartwood often with gum-like substances. Sometimes with laticifers containing gutta between cortex and phloem. Sclerenchymatous idioblasts often present. Silica bodies sometimes present. Cristarque cells sometimes present. Calciumoxalate as prismatic, rhomboidal, cuboid, acicular or irregular crystals, crystal sand, druses, raphides, or styloids. Calciumcarbonate and/or calciumsulfate sometimes present.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, furcate, T-shaped, malpighiaceous hairs, stellate, fasciculate, candelabra-shaped, bristle-like, prickly hairs, dendritic, lepidote or peltate (sometimes lanate, rarely vesicular), or absent; glands and glandular hairs sometimes present (rarely lepidote glands, helical glands or pearl glands); stinging hairs occasionally present.

Leaves Alternate or opposite (rarely verticillate), pinnately or palmately compound or simple and entire or lobed, with conduplicate, supervolute, convolute, involute, revolute, plicate, curved, circinate or flat ptyxis (rarely absent). Stipules interpetiolar, intrapetiolar, petiolar or cauline, free or connate, often rudimentary or absent (sometimes modified into spines, hairs or glands); leaf sheath absent. Colleters sometimes present. Venation pinnate or palmate, eucamptodromous, brochidodromous, craspedodromous, semicraspedodromous or reticulodromous (rarely acrodromous, actinodromous, campylodromous or parallelodromous). Stomata anomocytic, paracytic or anisocytic (rarely cyclocytic, laterocytic, helicocytic, staurocytic, hemiparacytic, diacytic, tetracytic or polycytic). Cuticular wax crystalloids as rodlets, platelets, scales, tubuli or rosettes of platelets (Fabales type) (sometimes absent). Domatia as pits, pockets or hair tufts, or absent. Epidermis with or without mucilaginous idioblasts, often containing druses; often with idioblasts (sometimes gland-dots and schizogenous secretory cavities) containing ethereal oils; epidermal cells sometimes with crystals. Lamina occasionally with resinous glands (colleters). Mesophyll with or without secretory cavities and/or ducts, with or without sclerenchymatous idioblasts, often with crystalliferous cells containing calciumoxalate as rhomboidal crystals or druses (sometimes styloids or crystal sand), or with sclerenchymatous idioblasts containing dendrosclereids or other types of sclereids; sometimes with tanniniferous cells. Laticifers with gutta sometimes present. Stomatal (including guard cells) or idioblastic myrosin cells (with myrosinase) sometimes frequent. Leaf margin or leaflet margins entire, crenate, sinuate or serrate; teeth with one vein proceeding into congested caducous tooth apex; glandular teeth rarely present on leaf margin. Leaf teeth sometimes begonioid, cucurbioid, cunonioid, malvoid, rosoid, salicoid, urticoid or violoid.

Inflorescence Terminal or axillary, panicle, fascicle, thyrse, thyrsoid, botryoid, bostrycoid, corymb, raceme-, spike-, catkin- or umbel-like cymose, or raceme, spike or catkin (sometimes pseudanthium). Bracts and/or floral prophylls (bracteoles) sometimes absent.

Flowers Actinomorphic or zygomorphic (rarely asymmetrical). Pedicel often articulated. Hypanthium sometimes present. Usually hypogyny (sometimes epigyny or half epigyny). Receptacle sometimes elongated into androgynophore or gynophore. Sepals (two to) four to five (to c. 20), usually with imbricate or imbricate-quincuncial, valvate or open (sometimes truncate, contorted or decussate, rarely induplicate-valvate or cochlear) aestivation, usually whorled (rarely [secondarily] spiral and indistinctly separate from petals), usually free (sometimes connate at base). Sepals often with three leaf traces from three gaps. Petals (two to) four or five (to more than 15), usually whorled (rarely spiral and indistinctly separate from sepals), with imbricate or imbricate-quincuncial, valvate, contorted or involute (sometimes crumpled, decussate or cochlear-descending, rarely cochlear, plicate or open) aestivation, usually free (sometimes more or less connate, rarely connate into campanulate corolla), or absent. Nectaries receptacular, supplied from receptacular or androecial traces, or on filament bases, or staminodial, or extrastaminal or intrastaminal, annular, cupular, unilateral or lobate, nectariferous disc, or as nectariferous glands of various shape, inserted on disc, perianth, adaxial side of hypanthium or as nectariferous hairs, or nectary absent. Disc present or absent. Flower often with typical mucilage cells having strongly thickened mucilaginous inner periclinal cell wall and distinct cytoplasm.

Androecium Stamens one to numerous (often 4+4 or 5+5; rarely more than 1.000), usually in one or more whorls, sometimes in three to five alternisepalous or antesepalous fascicles, usually haplostemonous or diplostemonous (sometimes obdiplostemonous, rarely triplostemonous), centripetally or centrifugally developing. Filaments free from each other or more or less connate, usually free from tepals (sometimes adnate at base to petals, epipetalous). Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely monosporangiate, disporangiate or trisporangiate), usually introrse (sometimes extrorse or latrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum usually secretory (rarely amoeboid-periplasmodial). Staminodia present or absent.

Pollen grains Microsporogenesis usually simultaneous (rarely successive). Pollen grains (2–)3(–11)-colp(oid)ate, (2–)3(–33)-colpor(oid)ate or (2–)3(–4)- to polypor(or)ate (rarely syncolpate or inaperturate; sometimes heterocolpate with pseudocolpi alternating with apertures), usually shed as monads (sometimes tetrads, rarely polyads), bicellular or tricellular at dispersal. Exine with usually columellate (sometimes granular or intermediate) infratectum.

Gynoecium Pistil composed of (one or) two to ten (to 20) usually connate (often paracarpous, rarely pseudomonomerous, rarely [secondarily] free), usually whorled antepetalous (rarely [secondarily] spiral) carpels; when three then median carpel adaxial. Ovary usually superior (sometimes inferior or semi-inferior), unilocular to quinquelocular (to 20-locular), sometimes on gynophore or androgynophore. Style single, simple, often hollow, or stylodia two to five (to 18), usually free, or absent. Stigma one, capitate, lobate, spatulate or peltate, or stigmas two to five, punctate, capitate, truncate or adaxially decurrent (rarely flabellate, fimbriate, penicillate or almost petaloid or as commissural ridges down style), usually papillate, usually Dry (sometimes Wet) type. Pistillodium usually absent (male flowers sometimes with pistillodium).

Ovules Placentation axile, apical, basal, subbasal or parietal (sometimes intrusively parietal, lateral, basal-lateral, marginal or free-central; rarely laminar). Ovules one to more than 300 per carpel, anatropous, campylotropous or hemianatropous (sometimes orthotropous, amphitropous or anacampylotropous, rarely pleurotropous), ascending, horizontal or pendulous, apotropous or epitropous, usually bitegmic, tenuinucellar or crassinucellar. Micropyle bistomal or endostomal (sometimes exostomal). Funicular, placental or stylar obturator sometimes present. Archespore sometimes bicellular or multicellular. Nucellar cap or nucellar beak sometimes present. Megagametophyte usually monosporic, Polygonum type (sometimes disporic, Allium or Endymion type, or tetrasporic, Penaea, Drusa or Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes proliferating, sometimes absent. Endosperm development usually nuclear (rarely helobial). Endosperm haustoria chalazal (sometimes lateral) or absent. Embryogenesis usually solanad, onagrad or asterad (rarely piperad or caryophyllad).

Fruit A loculicidal and/or septicidal (rarely septifragal or denticidal) capsule, berry, drupe, nut, samara or schizocarp (divided into two to five nut-like, samaroid, baccate or drupaceous mericarps; rarely a pyxidium, a hesperidium, a secondary syncarp, or an assemblage of achenes or follicles).

Seeds Perisperm rarely developed. Endosperm copious to sparse, oily, sometimes starchy, or absent. Embryo large or small, straight, plicate or curved (rarely hook-shaped, spirally twisted or circinate), usually well differentiated (rarely absent), oily, with or without chlorophyll. Cotyledons usually two (rarely three or four).

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

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, etc.) and their glycosides, flavones, flavone-C-glycosides, flavone methyleters, flavanone glycosides, afzelechin, biflavonoids, 5-deoxyflavonoids, desoxyflavonoids with B-ring, biflavanoids, trihydroxyflavonoids, methylated flavonoids, flavonoid sulfates, isoflavonoids, acylated anthocyanins, cyanidin, cyanidin-3-glycoside, delphinidin, apigenin, monoterpenoids and their esters, diterpene and triterpene derivatives, toxic diterpene esters, bitter-tasting tetracyclic and pentacyclic triterpenes and other triterpenes, tetranor- and pentanortriterpenes, dihydrochalcones, dammaranes, dammarane triterpenoids, surianol, phorbole ester diterpenes, oleanolic acid derivatives, arjunolic acid derivatives, quinoid and/or phenolic sesquiterpenes, gutta (trans-1,4-polyisoprene), catechin, oleanolic acid derivatives, ellagic, methylated ellagic and gallic acids, non-hydrolyzable tannins, ellagitannins, galloyltannins, proanthocyanidins (prodelphinidins), p-coumaric acid, caffeic acid, chlorogenic acid, cinnamic acid derivatives, hydroxycinnamate derivatives, protoalkaloids, tropane (hygrolinic) alkaloids, peptide alkaloids, indole alkaloids, quinazoline alkaloids, polyhydroxyalkaloids, pyridine alkaloids, loline alkaloids, quinolizidine alkaloids, pyrrolidine alkaloids, piperidine alkaloids, pyrrolizidine alkaloids (also as aliphatic monocarboxylic esters), benzylisoquinoline alkaloids, sesquiterpene alkaloids and other alkaloids (securinine, phyllantine, phyllochrisine, etc.), d-norpseudoephedrine (cathine), acacetin-7-O-glycoside, acacetin-7-O-diglycoside, glucosinolates (mustard oil glycosides) derived from phenylalanine, tyrosine, methionine, valine, isoleucine and/or leucine, triterpene saponins, pentacyclic terpene saponins, tyrosine-, leucine- or phenylalanine-derived cyanogenic compounds, phenol glycosides (salicin, populin etc.), cyclopentenoid (cyclopentenylic) cyanogenic glycosides and/or cyclopentenylic fatty acids, cyclopentenoid cyanhydrin glycosides derived from non-protein amino acid 2-(2-cyclopentenyl)glycine, toxic bufadienolides, steroidal and triterpene saponins, xanthones, polyacetate-derived anthraquinones and arthroquinones, anthraquinones, bitter anthracenones, naphthoquinones, narcotic L(S)-(-)-α-aminopropiophenone (cathinone), phlobaphene, bergenin, benzoquinone rapanone, benzoquinones (embelin), pyranochromones, β-sitosterol and its glycoside, hentriacontane, mesoinosite, maytansine (ansamycin macrolide), hexitols (dulcitol etc.), salacinol, cyclic polyvalent alcohols, punicic acid, eleostearic acid (isomere of punicic acid), hypericin, pseudohypericin, arbutin, emodin derivatives, biemodyles and closely allied compounds, benzophenones, acetophenones, anthrones, naphthodianthrones, coumarins, furanocoumarins, p-coumaric acid, coumarin derivatives substituted at position 4, syringaresinol, ferulic acid, phytosterols, quebrachitol, barbeyol, cannabinoids, humulones, lupulones, naphthalene and lupeolin derivatives, ethereal oils, hyperforin, picrotoxans, hydroxyproline betaines, mustard oils based on glucotropaeolin, glucocapparin, glucocleomin, citrullin, stilbenes, cholinesters etc., erucic acid, n-eicose-11-enoic acid, docosadienoic acid and other unsaturated fatty acids, sinapic acid, tartaric acid, tariric acid, tartric acid, lipids of cyclopropane and cyclopropenoid fatty acids and their derivatives, benzyl- and methoxybenzylisothiocyanates, myo-inositol, lignans, neolignans, nor-neolignans, pinitol, apiitol, N-methyltyrosine (non-protein amino acid), cyclic polyvalent alcohols, amides, type C18:3 fatty acids, cyclopropane amino acids, polygalitol, sweet-tasting proteins (brazzein, pentadin), and aromatic m-carboxycinnamic acids and proteolytic enzymes (papain, carpain), and nigracin present.

Fossils The oldest known flower fossil showing rosid affinities (with distinct calyx and corolla) emanate from the Late Albian to the Early Cenomanian of Nebraska. This ‘Rose Creek flower’ is pentamerous and hypogynous, with four series of floral parts, free persistent sepals and free thin petals, probably five antepetalous stamens, tricolporate pollen grains with psilate exine, and five connate carpels with free styles.

[Fabidae+Malvidae]

Potential synapomorphies: Mucilage cells sometimes with thickened inner periclinal walls and distinct cytoplasm. Endosperm scanty. Embryo elongated. Genome duplication. Plastid gene infA usually lost/non-functional (functional in Celastrales). Mitochondrial intron coxII.i3 lost. Plastid gene rps16 often entirely or partially lost. – This clade embraces all Rosidae except Vitaceae.

Rosidae have principally receptacular nectaries, whereas Vitaceae have gynoecial nectaries. The corolla often has a delayed development compared to other floral parts.

Phylogeny of Rosidae based on DNA data (Wang & al. 2009; Worberg & al. 2009; Soltis & al. 2011; somewhat modified). The bootstrap support for Rosidae was 85% (Soltis & al. 2011), whereas the support for the clades [Geraniales+Myrtales] and [Cucurbitales+Juglandales] was 79% and 76%, respectively. Vitaceae are sister to all other Rosidae (bootstrap support 100%, although 72% in Wang & al. 2009). Rosidae sensu stricto are split into the two clades Fabidae (Zygophyllales to Juglandales) and Malvidae (Myrtales to Capparales, BS support 97%). The branching pattern within these clades are highly supported (75% to 100%). The support for the COM clade (Celastrales, Oxalidales and Malpighiales) as sister to the Nitrogen-fixing clade (Polygalales, Rosales, Cucurbitales and Juglandales) was only 57%. The clade [Malpighiales+Oxalidales], to which Celastrales are sister-group, is supported by 59%. Zygophyllales may be sister to [COM+Nitrogen-fixing-clade], but the support for this is low. The COM clade is sister to Malvidae in a study by Qiu & al. 2010.

Fabidae W. S. Judd, D. E. Soltis et P. S. Soltis in Taxon 56: E29. Aug 2007 (eurosids I)

Potential synapomorphies: Extrafloral nectaries often consisting of palisade epidermal cells. Endosperm scanty. – A trichotomy is formed by Zygophyllales, the COM clade and the Nitrogen-fixing clade, or Zygophyllales is sister-group to a clade comprising the COM and the Nitrogen-fixing clades.

The COM clade

[Celastrales+[Oxalidales+Malpighiales]]

Potential synapomorphies: Ovules incompletely tenuinucellar. Seed coat exotegmic; exotegmen often fibrous of special type.

[Oxalidales+Malpighiales]

Potential synapomorphies: Inner integument often thicker than outer integument. Plastid gene infA lost or non-functioning. – The megasporangium is often thin and the seeds often arillate.

The Nitrogen-fixing clade

[Polygalales+[Rosales+[Cucurbitales+Juglandales]]]

Potential synapomorphies: Pre-adaptation to fixation of nitrogen by root-dwelling associates, usually actinobacteria of the genus Frankia or rhizobia (with exception in Cannabaceae). Tension wood present. Seed coat exotestal. Endosperm at most sparce. Embryo large.

Symbiosis with nitrogen-fixing Gram-positive actinobacteria, Frankia, seems to have evolved at least four times (possibly six times). The three lineages of Frankia have obviously diverged prior to the origin of angiosperms. Frankia strains are responsible for nitrogen-fixation in Cucurbitales (i.a. Coriaria and Datisca), Juglandales (e.g. Alnus, Casuarinaceae and Myricaceae) and Rosales (i.a. Elaeagnaceae and Rhamnaceae), whereas N-fixation is carried out by Gram-negative α-proteobacteria (Rhizobium) in Polygalales (some Mimosoideae and the majority of Faboideae inFabaceae) and in Parasponia (Cannabaceae in Rosales). β-proteobacteria are responsible for the nodule formation and the N-fixation in some Fabaceae clades.

The underlying molecular causes of developing nodular nitrogen-fixation are being explored and it seems that plasmid-borne genetic components, which may be exchanged between bacteria, are partly involved. Furthermore, many genes directing the development of vesicular-arbuscular mycorrhiza are identical to those involved in nodular N-fixation. Ectomycorrhiza is frequently occurring among groups in the Nitrogen-fixing clade. Nodulation probably involves interactions between several genes.

The Frankia nodules (and also the rhizobial nodules in Parasponia) form through the modification of lateral roots, a process directed by genes in the host plant. The pericyclic nodule initiation results in the establishment of N-fixing bacterial colonies in mid-cortical cells surrounding a central vascular cylinder. The nodule tissues are enclosed by an outer periderm. Intercellular penetration of root epidermis occurs in Rosales and may be a plesiomorphy, whereas in Cucurbitales and Juglandales the infection takes place via rhizoids (root hairs).

In Fabaceae the nodules develop from root cortical cell divisions and their vascular tissue becomes peripheral. The nodular initiation may take place by intercellular penetration, infection through rhizoids or infection at wound sites. The symbiosis between Rhizobium and the plant is a prerequisite of the nitrogen fixation, the fabaceous plant producing at least one of the co-factor components essential for the nitrogenase activity. The nitrogenase is present in Frankia in special symbiotic vesicles located on short lateral branches of the filamentous vegetative mycelium-forming bacterial cells. The vesicles differ in their morphology among the lineages of the Nitrogen-fixing clade (the NF clade).

Multiple parallel gains and losses have probably taken place during the evolution of nodular nitrogen-fixation among angiosperms. The majority of groups included in the NF clade seem to lack the capacity of forming root nodules and fix nitrogen. There may be a genetically based predisposition with a single origin for nodular N-fixation in the NF clade. If so, the common ancestor of the NF clade may have evolved the underlying genetic requirements for bacterial nodular N-fixation. Subsequently, parallel lineages would have the necessary genetic equipment for developing N-fixing mutualism. Alternatively, ancestors of many lineages in the NF clade may have lost the ability to develop the symbiosis.

Although there is high support for a single origin of nitrogen-fixing symbiosis, there are significant differences in nodule morphology among the four lineages of the NF clade. Apart from the rhizobial mutualism carried out by Gram-negative proteobacteria in Fabaceae, each of the remaining three (Frankia-inhabited) clades exhibits a distinct nodule type.

In Rosaceae, Elaeagnaceae and Rhamnaceae (Rosales) the infection is carried out by hyphal penetration of the middle lamella between epidermal cells of the root. Subsequent intercellular bacterial growth takes place towards the cortex of the root and, finally, penetration by the hyphae of cortical cells in a nodule primordium.

Coriariaceae and Datiscaceae (Cucurbitales) are infected via rhizoids (root hairs). They also have a specific arrangement of vesicles in infected cells, with vesicles orientated at right angles to a central vacuole instead of vesicles surrounding the periphery of the cytoplasm. Coriaria and Datisca possess well-aerated nodule tissue and their nodules have a specific cell layer that limits the diffusion of oxygen. Datisca produces nodule roots, as is the case in Myricaceae (Juglandales), whereas Coriaria does not produce nodule roots. Instead, Coriaria has lenticels, through which oxygen may pass into the nodule. Moreover, the non-infected cells are separate from the multinucleate infected cells.

The nodules of Casuarinaceae and Myricaceae (Juglandales) possess a specialized cell layer which has thick or lignified walls and is resistant against oxygen diffusion, resulting in a lower oxygen concentration in the Frankia-infected cells. The growth of the nodule lobes (formation of the nodular roots) in Juglandales is usually indeterminate. This probably results in higher absorption of oxygen due to the increased surface area. Alnus, which does not produce nodule roots, is an exception to this. Instead, Alnus (having well-aerated nodule tissue) possesses nodule lenticels, which are similar to those in Coriaria (Cucurbitales) and enhance the transport of free oxygen into the nodules (oxygen diffusion resistance layer is absent in Alnus). Furthermore, the haemoglobin concentration is usually higher in the nodules of Juglandales (again, Alnus is an exception).

[Rosales+[Cucurbitales+Juglandales]]

Potential synapomorphies: Placentation apical. Ovules one or two per carpel.

[Cucurbitales+Juglandales]

Potential synapomorphies: Epigyny (ovary inferior). Ovules two per carpel. Fruit single-seeded, indehiscent. – The gynoecium is usually tricarpellate.

Malvidae W. S. Judd, D. E. Soltis et P. S. Soltis in Taxon 56: E29. Aug 2007 (eurosids II)

Potential synapomorphies: Contorted petal aestivation. Presence of gynophore and/or androphore. Campylotropous ovules. Micropyle Z-shaped (zig-zag).

[Geraniales+Myrtales]

Potential synapomorphies: Calyx persistent in fruit. Ellagic acid present.

[Picramniaceae+[Sapindales+[Huerteales+[Malvales+Capparales]]]]

Potential synapomorphies: Placentation apical. Ovules two per carpel, pendulous. – This clade is sister to Staphyleales.

The relationships in Malvidae are unclear and it is not at all certain which group is sister to Capparales. The clade [Malvales+Sapindales] is a possible candidate, and the clade [Malvales+Capparales] has large support in some analyses, although the sampling was insufficient and Huerteales was not included.

[Sapindales+[Huerteales+[Malvales+Capparales]]]

Potential synapomorphies: Cambium sometimes storied. Vessel elements with simple perforation plates. Petiole vascular bundle transection annular. Style present. Inner integument thicker than outer. Endosperm scarce. Flavonols present.


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