Belongs within: Pan-Angiospermae.
Contains: Nymphaeales, Austrobaileyales, Mesangiospermae.
The Angiospermae are the flowering plants, one of the most speciose clades of plants alive today. Flowers possess a closed carpel that develops into a fruit, and ovules possess two integuments. Seeds ancestrally develop an endosperm via double fertilisation (Cantino et al. 2007). Cantino et al. (2007) defined the angiosperms as a crown clade (specifically, the most inclusive crown clade containing Zea mays but not Cycas, Ginkgo, Gnetum or Pinus).
Within the angiosperms, recent phylogenies are in agreement that the so-called ‘ANITA grade’ (named as an acronym for its component taxa: Amborella, Nymphaeales, Illicium, Trimenia, Austrobaileya) contains the outgroup(s) to the remaining angiosperms (dubbed the Mesangiospermae by Cantino et al. 2007). Within the ANITA grade, the Austrobaileyales (the -ITA) are most often identified as the direct sister group to the mesangiosperms, with Nymphaeales and Amborella as successively more distant sister groups. Amborella trichopoda is a small sprawling understorey shrub endemic to New Caledonia, previously classified with the Laurales before its recognition as the basalmost living angiosperm. It has apparently two-ranked exstipulate leaves with serrated or rippled margins. Plants are dioecious with small flowers with an undifferentiated perianth. Fruits are red drupes with a papery skin. Amborella trichopoda is distinguished from many other angiosperms in that its xylem lacks vessels, cells with perforated ends allowing for direct transmission of fluids between cells, and contains only tracheids, cells with pits in the end walls. However, vessels are also absent in Hydatellaceae, and the vessel morphology of Nymphaeales differs from that of Mesangiospermae and Austrobaileyales, so it remains uncertain when and how often vessels originated in flowering plants (Angiosperm Phylogeny Website).
Angiosperms that remain of uncertain affinities include Atrichodendron, a Vietnamese rainforest shrub historically placed in the Solanaceae.
The earliest fossils of likely crown-group angiosperms are known from the early Cretaceous with angiosperms becoming a dominant part of the flora in the latter part of that period. Fossil leaves attributed to the holding genus Phyllites may indicate an earlier origin for angiosperms but are not definitive. Other fossils of uncertain affinities include ‘Doliostrobus’ rerollei, originally described as a conifer (Kvaček 2002).
Synapomorphies (Cantino et al. 2007): Closed carpel developing into a fruit; ovule with two integuments; female gametophyte highly reduced, usually with no more than eight nuclei; endosperm resulting from double fertilisation; microgametophyte with three nuclei; two or more orders of leaf venation; axially aligned companion cells derived from same mother cells as sieve elements; pollen with unlaminated endexine; stamen with two pairs of pollen sacs.
<==Angiospermae (see below for synonymy)
|--AmborellaSD11 [Amborellaceae, AmborellalesAPG16]
| `--A. trichopoda Baill. 1869CD07
`--+--NymphaealesAPG16
`--+--AustrobaileyalesAPG16
`--MesangiospermaeAPG16
Angiospermae incertae sedis:
Kagenkia oblongaBS01
×Citrofortunella microcarpaC-R02
DicotilophyllumGT02
|--D. altenburgenseJD05
|--D. pusillumZ02
`--D. spatulatiumGT02
PhyllitesGT02
|--P. eucalyptroidesH79
|--P. platanoidesGT02
`--P. proteaceus Bozzi 1891GT02
Carcophyllum leogianenseGT02
Mancuna deeringianaKH02
Helecia serrataK03
Schindapsus perakensisK03
Pentastemona [Pentastemonaceae]H03
AppomattoxiaT00
Cynocrambe [Cynocrambaceae]YY22
Pileus conicaRN72b
Osmaronia dioicaGM96
‘Doliostrobus’ rerollei Marion 1884K02
Tristellateia australasiaeP88
Mira undulata Colenso 1845C45
AfropollisI02
|--A. jardinus (Brenner) Doyle et al. 1982I02
`--A. kahramanensis Ibrahim & Schrank 1995I02
Diporocolpopollenites Yi & Batten 2002YB02
`--*D. kachiensis Yi & Batten 2002YB02
Dilwynites Harris 1965YB02
`--*D. granulatus Harris 1965YB02
Bratzevaea amurensis (Bratzeva) Takahashi in Takahashi & Shimono 1982YB02
CallistopollenitesYB02
|--C. comis Srivastava 1969YB02
`--C. tumidoporus Srivastava 1969YB02
CaryapollenitesYB02
|--C. scabratus Groot & Groot 1962YB02
`--C. triangulus (Pflug) Krutzsch 1961YB02
CranwelliaYB02
|--C. edmontonensis Srivastava 1967YB02
`--C. striata (Couper) Srivastava 1967YB02
Fibulapollis mirificus (Chlonova) Chlonova 1961YB02
JianghanpollisYB02
|--J. mikros Wang & Zhao 1979YB02
`--J. radiatus Wang & Zhao 1979YB02
Leptopecopites pocockii (Srivastava) Srivastava 1978YB02
Penetetrapites inconspicuus Sweet 1986YB02
Porocolpopollenites vestibulum (Potonié) Thomson & Pflug 1953YB02
Retitrescolpites Sah 1967YB02
Tricolpites Cookson ex Couper 1953YB02
|--T. brevicolpusV12
|--T. lillieiV12
|--T. microreticulatus Belsky, Boltenhagen & Potonié 1965CP02
|--T. parvus Stanley 1965I02
|--T. reticulatus Cookson 1947YB02
`--T. vulgaris (Pierce) Srivastava 1969I02
Tricolporites prolata Cookson 1947YB02
Wodehouseia spinata Stanley 1961YB02
Brenneripollis peroreticulatus (Brenner) Juhász & Góczán 1985I02
CretacaeiporitesI02
|--C. densimurus Schrank & Ibrahim 1995I02
`--C. polygonalisI02
Dichastopollenites ghazalatensis Ibrahim 1996I02
Integritetradites porosus Schrank & Mahmoud 2000I02
RouseaI02
|--R. delicipollis Srivastava 1977I02
`--R. miculipollis Srivastava 1975I02
Striatopollis trochuensis (Srivastava) Farabee & Canright 1986I02
Delinea indicaYZ02
Dolichoglottis scorzoneroidesW91
Rhoipites cryptoporus Srivastava 1972YB02
Strepsolon jamesoniiMM96
Acomastylis rossiiGS99
Cocconia sessifoliaRN72a
AtrichodendronAPG16
HiraniaAPG16
Chattacchloia italicaP09
Cyclobalanopsis glaucaLI89
BlighiaF11
|--B. sapidaF11
`--B. unijugataDP72
Loureira Cavanilles 1799C99
|--L. cuneifolia Cavanilles 1799C99
`--L. glandulosa Cavanilles 179C99
Omphalocarpum Beauvois 1800B00
Merendera bulbocodium Ramond 1801R01
Antholithes wellsi Brown 1956B56
Quoya cuneata [=Pityrodia cuneata]H18
PetalostemonC01 [=Petalostemum (l. c.)M68]
|--P. candidusC00
`--P. oligophyllusC01
LepachysC01
|--L. columnarisC01
`--L. tagetesC01
Loranthophyllum dubiumH79
Galeobdolon luteumR13
Archaeanthus linnenbergeriSBM11
Helonias bullataSBM11
Notaphoebe umbelliflora [incl. Mastixia cuneata Bl. 1850]M77
HibiscadelphusD81
|--H. bombycinasD81
|--H. distansW92
`--H. wilderianusD81
Calvaria majorD81
Cordeauxia edulisD81
Schwartzia simplexWL10
Bertholletia excelsaKSM06
Bolivaria [Bolivariaceae]G39
|--B. integrifoliaG39
`--B. trifidaG39
Cevallia sinuataC00
Parosela scopariaC00
BatschiaJ23
|--B. canescensJ23
`--B. longifloraJ23
Symphoria glomerataJ23
Cristaria coccinea [=Malva coccinea]J23
Feronia elephantumT-W89
Pierardia dulcisT-W89
PleiospilosJ54
|--P. bolusiiJ54
|--P. dekenaniJ54
`--P. simulansJ54
Glottiphyllum linguiformeJ54
Cucifera thebaicaM10
Congdonia pinetorumM68
Angiospermae [Angiospermophyta, Angiospermopsida, Annonanae, Annoniflorae, Anthophyta, Archichlamydeae, Centrolepideae, Chloranthineae, Dicotyledoneae, Dicotyledones, Epigynae, Incompletae, Magnolianae, Magnoliopsida, Nymphaeidae, Polypetalae, Thalamifloreae]
*Type species of generic name indicated
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