Specimen of Amborella trichopoda, photographed by Stan Shebs.

Belongs within: Pan-Angiospermae.
Contains: Nymphaeales, Austrobaileyales, Mesangiospermae.

The Angiospermae are the flowering plants, one of the most speciose clades of plants alive today. 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).

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. Earlier fossil leaves attributed to the holding genus Phyllites (used for fossil leaves of uncertain affinities) may indicate an earlier origin for angiosperms but are not definitive.

Synapomorphies (from 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)
    |--Amborella [Amborellaceae, Amborellales] WM14
    |    `--A. trichopoda Baill. 1869 CD07
    `--+--Nymphaeales WM14
       `--+--Austrobaileyales WM14
          `--Mesangiospermae WM14

Angiospermae incertae sedis:
  Tristerix tetrandrus BS01
  Kagenkia oblonga BS01
  ×Citrofortunella microcarpa C-R02
  Dicotilophyllum GT02
    |--D. altenburgense JD05
    |--D. pusillum Z02
    `--D. spatulatium GT02
  Phyllites GT02
    |--P. eucalyptroides H79
    |--P. platanoides GT02
    `--P. proteaceus Bozzi 1891 GT02
  Carcophyllum leogianense GT02
  Mancuna deeringiana KH02
  Adenocalymma paulistarum HH03
  Parapiptadenia excelsa HH03
  Forrestia griffithii K03
  Helecia serrata K03
  Schindapsus perakensis K03
  Pentastemona [Pentastemonaceae] H03
  Appomattoxia T00
  Zippelia T00
  Cynocrambe [Cynocrambaceae] YY22
  Pileus conica RN72b
  Osmaronia dioica GM96
  ‘Doliostrobus’ rerollei Marion 1884 K02
  Tristellateia australasiae P88
  Mira undulata Colenso 1845 C45
  Afropollis I02
    |--A. jardinus (Brenner) Doyle et al. 1982 I02
    `--A. kahramanensis Ibrahim & Schrank 1995 I02
  Diporocolpopollenites Yi & Batten 2002 YB02
    `--*D. kachiensis Yi & Batten 2002 YB02
  Dilwynites Harris 1965 YB02
    `--*D. granulatus Harris 1965 YB02
  Bratzevaea amurensis (Bratzeva) Takahashi in Takahashi & Shimono 1982 YB02
  Callistopollenites YB02
    |--C. comis Srivastava 1969 YB02
    `--C. tumidoporus Srivastava 1969 YB02
  Caryapollenites YB02
    |--C. scabratus Groot & Groot 1962 YB02
    `--C. triangulus (Pflug) Krutzsch 1961 YB02
  Cranwellia YB02
    |--C. edmontonensis Srivastava 1967 YB02
    `--C. striata (Couper) Srivastava 1967 YB02
  Fibulapollis mirificus (Chlonova) Chlonova 1961 YB02
  Jianghanpollis YB02
    |--J. mikros Wang & Zhao 1979 YB02
    `--J. radiatus Wang & Zhao 1979 YB02
  Leptopecopites pocockii (Srivastava) Srivastava 1978 YB02
  Penetetrapites inconspicuus Sweet 1986 YB02
  Porocolpopollenites vestibulum (Potonié) Thomson & Pflug 1953 YB02
  Retitrescolpites Sah 1967 YB02
  Tricolpites Cookson ex Couper 1953 YB02
    |--T. brevicolpus V12
    |--T. lilliei V12
    |--T. microreticulatus Belsky, Boltenhagen & Potonié 1965 CP02
    |--T. parvus Stanley 1965 I02
    |--T. reticulatus Cookson 1947 YB02
    `--T. vulgaris (Pierce) Srivastava 1969 I02
  Tricolporites prolata Cookson 1947 YB02
  Wodehouseia spinata Stanley 1961 YB02
  Brenneripollis peroreticulatus (Brenner) Juhász & Góczán 1985 I02
  Cretacaeiporites I02
    |--C. densimurus Schrank & Ibrahim 1995 I02
    `--C. polygonalis I02
  Dichastopollenites ghazalatensis Ibrahim 1996 I02
  Integritetradites porosus Schrank & Mahmoud 2000 I02
  Rousea I02
    |--R. delicipollis Srivastava 1977 I02
    `--R. miculipollis Srivastava 1975 I02
  Striatopollis trochuensis (Srivastava) Farabee & Canright 1986 I02
  Delinea indica YZ02
  Payena lucida CK06
  Dolichoglottis scorzoneroides W91
  Rhoipites cryptoporus Srivastava 1972 YB02
  Strepsolon jamesonii MM96
  Acomastylis rossii GS99
  Cocconia sessifolia RN72a
  Atrichodendron APG16
  Hirania APG16
  Chattacchloia italica P09
  Cyclobalanopsis glauca LI89
  Fasanus acuminatus R35
  Blighia F11
    |--B. sapida F11
    `--B. unijugata DP72
  Loureira Cavanilles 1799 C99
    |--L. cuneifolia Cavanilles 1799 C99
    `--L. glandulosa Cavanilles 179 C99
  Omphalocarpum Beauvois 1800 B00
  Merendera bulbocodium Ramond 1801 R01
  Antholithes wellsi Brown 1956 B56
  Malocathrix glabrata S38
  Cyanella alba LY82
  Quoya cuneata [=Pityrodia cuneata] H18
  Petalostemon oligophyllus C01
  Lepachys C01
    |--L. columnaris C01
    `--L. tagetes C01
  Loranthophyllum dubium H79
  Galeobdolon luteum R13
  Jeffersonia diphylla SBM11
  Archaeanthus linnenbergeri SBM11
  Helonias bullata SBM11
  Notaphoebe umbelliflora [incl. Mastixia cuneata Bl. 1850] M77
  Hibiscadelphus D81
    |--H. bombycinas D81
    |--H. distans W92
    `--H. wilderianus D81
  Calvaria major D81
  Cordeauxia edulis D81
  Schwartzia simplex WL10
  Bertholletia excelsa KSM06
  Bolivaria [Bolivariaceae] G39
    |--B. integrifolia G39
    `--B. trifida G39

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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Last updated: 27 March 2021.

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