Specimen of Amborella trichopoda, photographed by Stan Shebs.

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

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, and is dioecious with small flowers with an undifferentiated perianth. 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).

Recent studies have also identified the Hydatellaceae as closely related to Nymphaeales within the ANITA grade, though they had previously been treated as monocotyledons (Saarela et al. 2007). Hydatellaceae are small, superficially grass-like aquatic plants that only reach a few centimetres in height. They have been classified into two genera based on whether individual flower-heads bear both male and female flowers (Trithuria), or male and female flowers are born on separate heads (Hydatella); this distinction is not universally recognised.

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] SR07
    |    `--A. trichopoda Baill. 1869 CD07
    `--+--+--Austrobaileyales SR07
       |  `--Mesangiospermae SR07
       `--+--Nymphaeales SR07
          `--Hydatellaceae [Hydatellales, Hydatellanae] SR07
               |--Hydatella Diels 1904 ME70
               |    |--*H. australis ME70
               |    `--H. inconspicua (Cheesem.) Cheesem. 1907 [=Trithuria inconspicua Cheesem. 1906] ME70
               `--Trithuria SR07 [incl. Juncella Mueller ex Hieronymus 1888 ME70]
                    |--T. bibracteata GK00
                    `--T. submersa SR07

Angiospermae incertae sedis:
  Tristerix tetrandrus BS01
  Kagenkia oblonga BS01
  Acroptilon repens BC01
  Santolina marchii R-CT01
  ×Citrofortunella microcarpa C-R02
  Dicotilophyllum GT02
    |--D. altenburgense JD05
    |--D. pusillum Z02
    `--D. spatulatium GT02
  Phyllites GT02
    |--P. platanoides GT02
    `--P. proteaceus Bozzi 1891 GT02
  Carcophyllum leogianense GT02
  Mancuna deeringiana KH02
  Adenocalymma paulistarum HH03
  Dunalia HH03
    |--D. breviflora HH03
    `--D. lycioides HH03
  Parapiptadenia excelsa HH03
  Forrestia griffithii K03
  Helecia serrata K03
  Micromelum minatum K03
  Parashorea malaanonan K03
  Schindapsus perakensis K03
  Pentastemona [Pentastemonaceae] H03
  Appomattoxia T00
  Codon T00
  Zippelia T00
  Corynocarpus Forster & Forster 1776 [Corynocarpaceae] A61
    |--C. cribbianus (Bailey) Smith 1956 (see below for synonymy) F78
    `--C. laevigatus Forster & Forster 1776 A61
  Haptanthus T00
  Heteranthia T00
  Pteleocarpa T00
  Cynocrambe [Cynocrambaceae] YY22
  Pileus conica RN72
  Plectritis congesta GM96
  Osmaronia dioica GM96
  ‘Doliostrobus’ rerollei Marion 1884 K02
  Pemphis acidula P88
  Tristellateia australasiae P88
  Mira undulata Colenso 1845 C45
  Lesquerella fendleri KIW98
  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
  Retitricolpites anguloluminosus (Anderson) Yi & Batten 2002 (see below for synonymy) YB02
  Aquilapollenites YB02
  Alnipollenites YB02
    |--A. trina (Stanley) Norton in Norton & Hall 1969 YB02
    `--A. verus (Potonié) Potonié 1960 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
  Loranthacites catterallii Srivastava 1969 YB02
  Penetetrapites inconspicuus Sweet 1986 YB02
  Porocolpopollenites vestibulum (Potonié) Thomson & Pflug 1953 YB02
  Proteacidites YB02
    |--P. magnus Samoilovitch 1961 YB02
    |--P. retusus Anderson 1960 YB02
    `--P. thalmannii Anderson 1960 YB02
  Retitrescolpites Sah 1967 YB02
  Tricolpites Cookson ex Couper 1953 YB02
    |--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
  Retimonocolpites variplicatus Schrank & Mahmoud 1998 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
  Aegopodium podagraria HM07
  Rhoipites cryptoporus Srivastava 1972 YB02
  Strepsolon jamesonii MM96
  Macadamia integrifolia B88
  Lambertia inermis R96
  Acomastylis rossii GS99
  Chilopsis linearis BM76

Nomen nudum: Juncella tasmanica Mueller 1854 ME70

Angiospermae [Angiospermophyta, Angiospermopsida, Annonanae, Annoniflorae, Anthophyta, Archichlamydeae, Chloranthineae, Dicotyledoneae, Magnolianae, Magnoliopsida, Nymphaeidae]

Corynocarpus cribbianus (Bailey) Smith 1956 [=Cyanocarpus cribbiana Bailey 1897, Helicia cribbiana (Bailey) Bailey 1901; incl. Corynocarpus australasica White 1933] F78

Retitricolpites anguloluminosus (Anderson) Yi & Batten 2002 [=Tricolporites anguloluminosus Anderson 1960] YB02

*Type species of generic name indicated


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