Angiospermae

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The earliest fossil angiosperm, ''[[Archaefructus]]'', comes from the [[Yixian]] formation in China and is dated to about 125 million years BP (Sun ''et al.'', 2002). Angiosperm pollen has been found in the fossil record perhaps as long ago as 130 million years.
The earliest fossil angiosperm, ''[[Archaefructus]]'', comes from the [[Yixian]] formation in China and is dated to about 125 million years BP (Sun ''et al.'', 2002). Angiosperm pollen has been found in the fossil record perhaps as long ago as 130 million years.
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The relationships of angiosperms to other plant [[taxon|taxa]] remain contentious (see Friedman & Floyd, 2001, for an overview). Morphological data indicated that the [[Gymnospermae|gymnosperms]] were [[paraphyletic]] with regard to the angiosperms, and that the [[Gnetales]] were the closest living relatives of the angiosperms. However, molecular data have indicated that modern gymnosperms form a [[monophyletic]] sister group to angiosperms, with the Gnetales more closely related to (possibly even within) the [[Pinopsida|conifers]]. However, as the gymnosperm [[crown group]] dates back to the [[Carboniferous]], the angiosperm [[stem group|stem]] must have diverged by that time if the molecular analyses are correct. A number of fossil seed plant groups of uncertain relationships are known from within that time frame, and it seems likely that at least some of these taxa lie on the angiosperm stem. Taylor ''et al.'' (2006) demonstrated that [[oleanane]], a [[diagenesis|diagenetic]] product of organic compounds found in angiosperms but absent from living gymnosperms, was present in fossils of the [[Cretaceous]] [[Bennettitales]] and [[Permian]] [[Gigantopteridales]], making these two groups likely angiosperm stem candidates. Other fossil taxa that have been suggested as angiosperm relatives include the [[Glossopteridales]] and ''[[Caytonia]]''.
+
The relationships of angiosperms to other plant [[taxon|taxa]] remain contentious (see Friedman & Floyd, 2001, for an overview). Morphological data indicated that the [[Gymnospermae|gymnosperms]] were [[paraphyletic]] with regard to the angiosperms, and that the [[Gnetales]] were the closest living relatives of the angiosperms. However, molecular data have indicated that modern gymnosperms form a [[monophyletic]] sister group to angiosperms, with the Gnetales more closely related to (possibly even within) the [[Pinopsida|conifers]]. However, as the gymnosperm [[crown group]] dates back to the [[Carboniferous]], the angiosperm [[stem group|stem]] must have diverged by that time if the molecular analyses are correct. A number of fossil seed plant groups of uncertain relationships are known from within that time frame, and it seems likely that at least some of these taxa lie on the angiosperm stem. Taylor ''et al.'' (2003) demonstrated that [[oleanane]], a [[diagenesis|diagenetic]] product of organic compounds found in angiosperms but absent from living gymnosperms, was present in fossils of the [[Cretaceous]] [[Bennettitales]] and [[Permian]] [[Gigantopteridales]], making these two groups likely angiosperm stem candidates. Other fossil taxa that have been suggested as angiosperm relatives include the [[Glossopteridales]] and ''[[Caytonia]]''.
Within the angiosperms, recent phylogenetic analyses have mostly agreed that the so-called ANITA grade (including ''[[Amborella]]'', [[Nymphaeales]] and [[Austrobaileyales]]), includes the basalmost living clades. The Austrobaileyales are most likely closer to the remaining angiosperms than are ''Amborella'' and Nymphaeales. Analyses disagree on whether ''Amborella'' alone or an ''Amborella'' + Nymphaeales clade represents the basalmost branch of angiosperms, but the former option is perhaps the more popular. Saarela ''et al.'' (2007) recently demonstrated that the [[Hydatellaceae]] also fall in this area as the sister group to Nymphaeales. The other major angiosperm clades (listed below) form a monophyletic group, but relationships between the clades are uncertain.
Within the angiosperms, recent phylogenetic analyses have mostly agreed that the so-called ANITA grade (including ''[[Amborella]]'', [[Nymphaeales]] and [[Austrobaileyales]]), includes the basalmost living clades. The Austrobaileyales are most likely closer to the remaining angiosperms than are ''Amborella'' and Nymphaeales. Analyses disagree on whether ''Amborella'' alone or an ''Amborella'' + Nymphaeales clade represents the basalmost branch of angiosperms, but the former option is perhaps the more popular. Saarela ''et al.'' (2007) recently demonstrated that the [[Hydatellaceae]] also fall in this area as the sister group to Nymphaeales. The other major angiosperm clades (listed below) form a monophyletic group, but relationships between the clades are uncertain.
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In most taxonomies, the flowering plants are treated as a coherent group. The most popular descriptive name has been Angiospermae (Angiosperms), with Anthophyta ("flowering plants") a second choice. These names are not linked to any rank. The [[Wettstein system]] and the [[Engler system]] use the name Angiospermae, at the assigned rank of subdivision.  The [[Reveal system]] treated flowering plants as subdivision [[Magnoliophytina]] (Frohne & U. Jensen ''ex'' Reveal, Phytologia 79: 70 1996), but later split it to [[Magnoliopsida]], [[Liliopsida]] and [[Rosopsida]]. The [[Takhtajan system]] and [[Cronquist system]] treat this group at the rank of [[division]], leading to the name Magnoliophyta. The [[Dahlgren system]] and [[Thorne system]] treat this group at the rank of class, leading to the name Magnoliopsida. However, the [[APG system]] of 1998 and the [[APG II system]] of 2003 do not treat angiosperms as a formal taxon but rather an informal clade.
In most taxonomies, the flowering plants are treated as a coherent group. The most popular descriptive name has been Angiospermae (Angiosperms), with Anthophyta ("flowering plants") a second choice. These names are not linked to any rank. The [[Wettstein system]] and the [[Engler system]] use the name Angiospermae, at the assigned rank of subdivision.  The [[Reveal system]] treated flowering plants as subdivision [[Magnoliophytina]] (Frohne & U. Jensen ''ex'' Reveal, Phytologia 79: 70 1996), but later split it to [[Magnoliopsida]], [[Liliopsida]] and [[Rosopsida]]. The [[Takhtajan system]] and [[Cronquist system]] treat this group at the rank of [[division]], leading to the name Magnoliophyta. The [[Dahlgren system]] and [[Thorne system]] treat this group at the rank of class, leading to the name Magnoliopsida. However, the [[APG system]] of 1998 and the [[APG II system]] of 2003 do not treat angiosperms as a formal taxon but rather an informal clade.
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==Phylogeny==
 
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As noted above, relationships between the major angiosperm clades are uncertain, and the tree below represents just one of the topologies that have been suggested.
 
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[[Progymnospermida|<==]]Angiospermae (see below for synonymy)
 
-
    |--''Archaefructus'' [Archaefructaceae]
 
-
    |    |--*''A. liaoningensis''
 
-
    |    `--''A. sinensis'' Sun, Ji ''et al.'' 2002
 
-
    `--+--''Amborella'' [Amborellaceae]
 
-
      |    `--''A. trichopoda''
 
-
      `--+--+--[[Nymphaeales]]
 
-
          |  `--Hydatellaceae [Hydatellales, Hydatellanae]
 
-
          |      |--''Hydatella inconspicua''
 
-
          |      `--''Trithuria submersa''
 
-
          `--+--[[Austrobaileyales]]
 
-
            `--+--Chloranthaceae
 
-
                |    |  ''i. s.'': ''Chloranthistemon''
 
-
                |    |--''Hedyosmum''
 
-
                |    `--+--''Chloranthus multistachys''
 
-
                |      `--+--''Sarcandra''
 
-
                |          `--''Ascarina''
 
-
                `--+--+--+--[[Ranunculidae]]
 
-
                  |  |  `--''Ceratophyllum'' [Ceratophyllaceae, Ceratophyllales]
 
-
                  |  |      |--''C. demersum''
 
-
                  |  |      `--''C. submersum''
 
-
                  |  `--[[Monocotyledones]]
 
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                  `--+--+--[[Magnoliales]]
 
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                      |  `--[[Laurales]]
 
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                      `--+--+--[[Piperales]]
 
-
                        |  `--[[Aristolochiales]]
 
-
                        `--Winterineae
 
-
                              |--''Drimys'' [Winteraceae, Winterales]
 
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                              |    |--''D. piperita''
 
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                              |    `--''D. winteri''
 
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                              `--Canellales
 
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                                  |--''Canella'' [Canellaceae]
 
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                                  `--''Tasmannia''
 
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Angiospermae [Angiospermophyta, Annonanae, Anthophyta, Chloranthineae, Dicotyledoneae, Magnolianae, Magnoliidae]
 
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Angiospermae ''incertae sedis'':
 
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    ''Lycium''
 
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      |--''L. afrum''
 
-
      |--''L. barbarum''
 
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      |--''L. berlandieri''
 
-
      `--''L. ferocissimum''
 
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    ''Coffea''
 
-
      |--''C. arabica''
 
-
      |--''C. canephora''
 
-
      `--''C. liberica''
 
-
    ''Ileostylus micranthus''
 
-
    ''Alepis flavida''
 
-
    ''Peraxilla''
 
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      |--''P. colensoi''
 
-
      `--''P. tetrapetala''
 
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    ''Tristerix tetrandrus''
 
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    ''Kagenkia oblonga''
 
-
    ''Forsythia''
 
-
    ''Pseudobotrys''
 
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    ''Leucanthemum vulgare''
 
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    ''Hoheria angustafolia''
 
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    ''Chondrilla''
 
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      |--''C. juncea''
 
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      `--''C. prenanthoides''
 
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    ''Acroptilon repens''
 
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    ''Santolina marchii''
 
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    ''Cynara cardunculus''
 
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    ''Murraya''
 
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    ''Pometia''
 
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    ×''Citrofortunella microcarpa''
 
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    ''Beilschmiedia''
 
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      |--''B. taraire''
 
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      `--''B. tawa''
 
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    ''Trilepidea adamsii''
 
-
    ''Dicotilophyllum''
 
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      |--''D. pusillum''
 
-
      `--''D. spatulatium''
 
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    ''Agoseris cuspidata''
 
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    ''Muhlenbergia cuspidata''
 
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    ''Phyllites''
 
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      |--''P. platanoides''
 
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      `--''P. proteaceus'' Bozzi 1891
 
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    ''Carcophyllum leogianense''
 
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    ''Mancuna deeringiana''
 
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    ''[[Inga]]''
 
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    ''Erythrina crista-galli''
 
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    ''Salpichroa''
 
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      |--''S. diffusa''
 
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      |--''S. origanifolia''
 
-
      `--''S. rhomboidea''
 
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    ''Adenocalymma paulistarum''
 
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    ''Acnistus''
 
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      |--''A. arborescens''
 
-
      |--''A. breviflorus''
 
-
      `--''A. parviflorus''
 
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    ''Dunalia''
 
-
      |--''D. breviflora''
 
-
      `--''D. lycioides''
 
-
    ''Parapiptadenia excelsa''
 
-
    ''Strobilanthes kunthiana''
 
-
    ''Donax'' [Marantaceae]
 
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      `--''D. canniformis''
 
-
    ''Clausena excavata''
 
-
    ''Forrestia griffithii''
 
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    ''Helecia serrata''
 
-
    ''Micromelum minatum''
 
-
    ''Parashorea malaanonan''
 
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    ''Pronephrium asperum''
 
-
    ''Schindapsus perakensis''
 
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    ''Stenochlaena palustris''
 
-
    ''Urophyllum nigricans''
 
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    ''Trianthema turgidifolia''
 
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    ''Eriospermum'' [Eriospermaceae]
 
-
    ''Pentastemona'' [Pentastemonaceae]
 
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    ''Alternanthera''
 
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      |--''A. achyrantha''
 
-
      |--''A. denticulata''
 
-
      |--''A. philoxeroides''
 
-
      `--''A. sessilis''
 
-
    ''Appomattoxia''
 
-
    ''Codon''
 
-
    ''Zippelia''
 
-
    ''Berberidopsis'' [Berberidopsidaceae]
 
-
    ''Corynocarpus'' [Corynocarpaceae]
 
-
      `--''C. laevigatus''
 
-
    ''Haptanthus''
 
-
    ''Heteranthia''
 
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    ''Pteleocarpa''
 
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    Cynocrambaceae
 
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      |--''Cynocrambe''
 
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      `--''Theligonum cynocrambe''
 
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    ''Pileus conica''
 
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    ''Plectritis congesta''
 
-
    ''Osmaronia dioica''
 
-
    ''‘Doliostrobus’ rerollei'' Marion 1884
 
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    ''Pongamia pinnata''
 
-
    ''Pemphis acidula''
 
-
    ''Tristellateia australasiae''
 
-
    ''Thespesia''
 
-
      |--''T. grandiflora''
 
-
      `--''T. populnea''
 
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    ''Mira undulata'' Colenso 1845
 
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    ''Hippeastrum aulicum''
 
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    ''Lesquerella fendleri''
 
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    ''Ipomopsis aggregata''
 
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    ''Cyanella'' [Tecophileaceae]
 
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    ''Calochortus longebarbatus''
 
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    ''Ugni molinae''
 
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    ''Afropollis''
 
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      |--''A. jardinus'' (Brenner) Doyle ''et al.'' 1982
 
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      `--''A. kahramanensis'' Ibrahim & Schrank 1995
 
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    ''Diporocolpopollenites'' Yi & Batten 2002
 
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      `--*''D. kachiensis'' Yi & Batten 2002
 
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    ''Dilwynites'' Harris 1965
 
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      `--*''D. granulatus'' Harris 1965
 
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    ''Retitricolpites anguloluminosus'' (Anderson) Yi & Batten 2002 (see below for synonymy)
 
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    ''[[Aquilapollenites]]''
 
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    ''Alnipollenites''
 
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      |--''A. trina'' (Stanley) Norton ''in'' Norton & Hall 1969
 
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      `--''A. verus'' (Potonié) Potonié 1960
 
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    ''Bratzevaea amurensis'' (Bratzeva) Takahashi ''in'' Takahashi & Shimono 1982
 
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    ''Callistopollenites''
 
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      |--''C. comis'' Srivastava 1969
 
-
      `--''C. tumidoporus'' Srivastava 1969
 
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    ''Caryapollenites''
 
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      |--''C. scabratus'' Groot & Groot 1962
 
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      `--''C. triangulus'' (Pflug) Krutzsch 1961
 
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    ''Cranwellia''
 
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      |--''C. edmontonensis'' Srivastava 1967
 
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      `--''C. striata'' (Couper) Srivastava 1967
 
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    ''Fibulapollis mirificus'' (Chlonova) Chlonova 1961
 
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    ''Jianghanpollis''
 
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      |--''J. mikros'' Wang & Zhao 1979
 
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      `--''J. radiatus'' Wang & Zhao 1979
 
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    ''Leptopecopites pocockii'' (Srivastava) Srivastava 1978
 
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    ''Loranthacites catterallii'' Srivastava 1969
 
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    ''Penetetrapites inconspicuus'' Sweet 1986
 
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    ''Porocolpopollenites vestibulum'' (Potonié) Thomson & Pflug 1953
 
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    ''Proteacidites''
 
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      |--''P. magnus'' Samoilovitch 1961
 
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      |--''P. retusus'' Anderson 1960
 
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      `--''P. thalmannii'' Anderson 1960
 
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    ''Retitrescolpites'' Sah 1967
 
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    ''Tricolpites'' Cookson ''ex'' Couper 1953
 
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      |--''T. microreticulatus'' Belsky, Boltenhagen & Potonié 1965
 
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      |--''T. parvus'' Stanley 1965
 
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      |--''T. prolata'' Cookson 1947
 
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      `--''T. vulgaris'' (Pierce) Srivastava 1969
 
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    ''Tricolporites prolata'' Cookson 1947
 
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    ''Wodehouseia spinata'' Stanley 1961
 
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    ''Brenneripollis peroreticulatus'' (Brenner) Juhász & Góczán 1985
 
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    ''Cretacaeiporites''
 
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      |--''C. densimurus'' Schrank & Ibrahim 1995
 
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      |--''C. mulleri''
 
-
      `--''C. polygonalis''
 
-
    ''Dichastopollenites ghazalatensis'' Ibrahim 1996
 
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    ''Integritetradites porosus'' Schrank & Mahmoud 2000
 
-
    ''Retimonocolpites variplicatus'' Schrank & Mahmoud 1998
 
-
    ''Rousea''
 
-
      |--''R. delicipollis'' Srivastava 1977
 
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      `--''R. miculipollis'' Srivastava 1975
 
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    ''Striatopollis trochuensis'' (Srivastava) Farabee & Canright 1986
 
-
 
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''Retitricolpites anguloluminosus'' (Anderson) Yi & Batten 2002 [=''Tricolporites anguloluminosus'' Anderson 1960]
 
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<nowiki>*</nowiki> Type species of genus indicated
 
==Links==
==Links==
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