Streptophyta

Coleochaete sp., copyright Micropolitan.org.


Belongs within: Corticata.
Contains: Sycidiaceae, Umbellinaceae, Porocharaceae, Characeae, Zygnematophyceae, Bryata, Apo-Tracheophyta.

The Streptophyta are a clade uniting the land plants (Embryophyta) with their closest allies among the green algae. Members of the Coleochaetales, Characeae and Embryophyta are united by the presence of branching growth, oogamous sexual reproduction and phragmoplastic cell division. Conversely, molecular analyses indicate that the basal members of the streptophyte lineage include the unicellular scaly alga Mesostigma viride (Karol et al. 2001). Members of the Klebsormidiophyceae grow as coccoid cells or unbranched filaments.

The clade Embryophyta contains the land plants, the dominant producers of the terrestrial environment. These organisms were able to thrive out of the water due to the retention and fertilisation of their eggs within protective structures, though less derived forms remain dependent on damp conditions for successful fertilisation. Land plants are often divided for convenience between the vascular (tracheophytes) and non-vascular (mosses, liverworts and hornworts) plants. Past classifications have used the name 'Bryophyta' to refer to all non-vascular plants, but the majority of recent authors would restrict this name to the mosses alone. There has been much debate over the exact interrelationships between the vascular and non-vascular plants. Many authors place the hornworts (Anthocerotophyta) as sister to the tracheophytes, as supported by the relationship between sporophytes and gametophytes: liverworts and mosses have the sporophyte and gametophyte separated by a placental space, usually filled with collapsed cells derived from the gametophyte, while hornworts and basal tracheophytes have the two in direct contact without a separating layer of collapsed cells (Frey et al. 2001). However, a recent large-scale molecular analysis by Wickett et al. (2014) suggested the possibility of a monophyletic clade of non-vascular plants, or a sister relationship between Anthocerotophyta and all other land plants. Fossil spores indicate that the land plants first arose during the Ordovician, with the four major living lineages possibly diverging by the end of that period. However, plant macrofossils have not yet been recovered from earlier than the Late Silurian (Bateman et al. 1998). Despite their potentially nested position, vascular plants are better represented in the early fossil record than non-vascular plants due to their higher preservation potential; unequivocal liverworts are not known from early than the Late Devonian (Bateman et al. 1998).

Embryophytes are characterised by alternation in generations between asexually-reproducing diploid sporophytes and sexually-reproducing haploid gametophytes. In liverworts, mosses and hornworts, the gametophytes are the dominant generation, and support the nutritively dependent sporophytes. In vascular plants, the sporophytes are independently growing and the dominant life stage. The Early Devonian Horneophyton is a stem-vascular plant that possessed both independently growing sporophytes and gametophytes, but did not possess the reinforced water-conducting tracheid cells of vascular plants (instead, it possessed a plesiomorphic central strand of conducting tissue that was not reinforced by thickenings). Horneophyton and vascular plants also share the production of multiple sporangia per individual. Horneophyton had terminal sporangia with a central columella (sterile section) similar to the sporangium of a hornwort.

Koeppenia eifeliensis Schweitzer 2000 is a strobilus (reproductive stalk) of uncertain affinities from the Lower Devonian of Denmark, which bore a dense arrangement of sporangiophores along its length on narrow branching stalks. Other fossils listed below include Rhabdotarachnoides simoni, originally described as an arachnid from the Lower Permian of Germany but since re-interpreted as an unidentifiable piece of plant material (Dunlop 2007). Neocystites, originally described as a cystoid echinoderm, probably represents a fragment of root (Kesling 1967).

Characters (from Adl et al. 2012, as Charophyta): Asymmetric motile cells, when present, with pair of cilia without mastigonemes; basal bodies with distinctive multilayered structure of microtubular rootlet and cytoskeletal anchor; thylakoids stacked; plastid with two membranes without periplastid endoplasmic reticulum; starch inside plastid; open mitosis; usually with phycoplast, but some with phragmoplast and cell plate; with primary plasmodesmata between adjacent cells in filamentous forms; filaments branching or nonbranching; with nonmotile vegetative phase; some with multinucleate cells; with or without sexual reproduction; sexual species with haplobiontic life cycle; with desiccation-resistant cysts (zygospores); glycolate oxidase in peroxisomes; Cu/Zn superoxide dismutase; ciliary peroxisome.

<==Streptophyta (see below for synonymy) WM14
    |--+--Mesostigma Lauterborn 1894 WM14, AS12 [Mesostigmatales, Mesostigmatophyceae]
    |  |    `--M. viride KM01
    |  `--+--Chlorokybus Geitler 1942 WM14, AS12 [Chlorokybales, Chlorokybophyceae, Chlorokybophyta]
    |     |    `--C. atmophyticus SG99
    |     `--Spirotaenia WM14
    |          |--*S. condensata WM14
    |          `--S. minuta WM14
    `--+--Klebsormidiophyceae AS12
       |    |--Interfilum AS12
       |    |--Entransia Hughes 1943 AS12, MK00
       |    |    `--E. fimbriata KM01
       |    `--Klebsormidium [Klebsormidiales, Klebsormidiobiotes, Klebsormidiophyta] WM14
       |         |  i. s.: K. subtile WM14
       |         |--K. flaccidum KM01
       |         `--+--K. nitens KM01
       |            `--K. subtilissimum KM01
       `--Phragmoplastophyta AS12
            |--Characeae KM01
            `--+--Coleochaetales [Chaetophycidae, Coleochaetophyceae] WM14
               |    |--Chaetosphaeridium (see below for synonymy) KM01
               |    |    |--C. globosum KM01
               |    |    `--C. ovalis KM01
               |    `--Coleochaete [Coleochaetobiotes, Coleochaetophyta] KM01
               |         |  i. s.: C. scutata SG99
               |         |--+--C. irregularis KM01
               |         |  `--C. sieminskiana KM01
               |         `--+--C. orbicularis KM01
               |            `--C. soluta KM01
               `--+--Zygnematophyceae WM14
                  `--Embryophyta [Cormophyta, Cryptogamia, Embryobiotes, Hepatophyta] WM14
                       |  i. s.: Eophyllophyton bellum K01
                       |         Sublepidodendron ZA02
                       |           |--S. mirabile ZA02
                       |           `--S. wusihense [=Lepidodendron wusihense, Eolepidodendron wusihense] CWC87
                       |         Colophospermum mopane WS02
                       |         Neocalamites nathorsti HH09
                       |         Samanea saman PP03
                       |         Onychiopsis Yokoyama 1889 BO02
                       |           `--O. psilotoides BO02
                       |         Isoetites BO02
                       |           |--I. abundans Tosolini & McLoughlin 2002 TMD02
                       |           `--I. daharensis Barale 1999 BO02
                       |         Ptilophyllum VB02
                       |         Otozamites BO02
                       |         Eranthis hyemalis V02
                       |         Synsepalum dulcificum ZB03
                       |         Meum athamanticum HH03
                       |         Rhaciopteris busseana A38
                       |         Diplothmema subfurcatum A38
                       |         Megalopteris dawsoni A38
                       |         Sporangites acuminata A38
                       |         Pterispermostrobus bifurcatus A38
                       |         Whittleseya A38
                       |           |--W. concinna A38
                       |           `--W. dawsoniana A38
                       |         Cardiocarpon A38
                       |           |--C. baileyi A38
                       |           |--C. cornutum A38
                       |           |--C. crampi A38
                       |           `--C. obliquum A38
                       |         Asterophyllites A38
                       |           |--A. acicularis [incl. A. equisetiformis] A38
                       |           `--A. parvulus [incl. A. grandis] A38
                       |         Yuccites vogesiacus W70
                       |         Campsiandra laurifolia DS86
                       |         Copaifera reticulata DS86
                       |         Radiola linoides [=Linum radiola] C55a
                       |         Succowia balearica [=Bunias balearica] C55a
                       |         Dentaria C55a
                       |           |--D. bulbifera C55a
                       |           `--D. enneaphylla C55a
                       |         Aretia villosa C55a
                       |         Nauenburgia thyrsiflora C55a
                       |         Ditassa lanceolata C55a
                       |         Cybistax antisyphilitica C55a
                       |         Nonnea ventricosa C55a
                       |         Doratanthera linearis C55a
                       |         Ramphicarpa fistulosa C55a
                       |         Anodendron paniculatum C55a
                       |         Lachnopylis oppositifolia C55a
                       |         Schultesia C55a
                       |           |--S. brachyptera C55a
                       |           `--S. stenophylla C55b
                       |         Voyria uniflora C55a
                       |         Micranthemum orbiculatum C55a
                       |         Hugueninia tanacetifolia C55a
                       |         Adenostyles leucophylla C55a
                       |         Kernera saxatilis C55a
                       |         Petrocallis pyrenaica C55a
                       |         Orobus luteus C55a
                       |         Athamantha C55a
                       |           |--A. cretensis C55a
                       |           `--A. libanotis C55b
                       |         Libanotis montana C55a
                       |         Serratula nudicaulis C55a
                       |         Betonica alopecurus C55a
                       |         Zephyritis taitensis C55a
                       |         Catabrosa aquatica [=Aira aquatica] C55a
                       |         Tristegis glutinosa C55b
                       |         Crambe C55b
                       |           |--C. hispanica C55b
                       |           |--C. maritima C55a
                       |           `--C. orientalis C55b
                       |         Cucubalus bacciferus C55b
                       |         Choerophyllum C55b
                       |           |--C. aromaticum C55b
                       |           `--C. aureum C55b
                       |         Imperatoria ostruthium [=Peucedanum ostruthium] C55b
                       |         Pyrethrum parthenium C55b
                       |         Leonurus C55b
                       |           |--L. cardiaca C55b
                       |           `--L. sibiricus C55b
                       |         Hyssopus officinalis C55b
                       |         Blitum C55b
                       |           |--B. polymorphum C55b
                       |           `--B. virgatum C55b
                       |         Anacharis alsinastrum C55b
                       |         Ervum C55b
                       |           |--E. hirsutum C55b
                       |           `--E. lens C55b
                       |         Helosciadum C55b
                       |           |--H. leptophyllum C55a
                       |           `--H. nodiflorum C55b
                       |         Lappa major C55b
                       |         Ambrina D03
                       |           |--A. ambrosioides D03
                       |           `--A. botrys C55b
                       |         Agathophytum bonus-henricus [=Blitum bonus-henricus] C55b
                       |         Guilandina bonduc [incl. G. bonducella] C55b
                       |         Drepanocarpus lunatus C55b
                       |         Ecastaphyllum brownei C55b
                       |         Schwenkia americana C55b
                       |         Capraria biflora C55b
                       |         Telanthera C55b
                       |           |--T. brasiliana C55b
                       |           |--T. frutescens C55b
                       |           `--T. maritima C55b
                       |         Remirea maritima C55b
                       |         Poinciana pulcherrima C55b
                       |         Ptarmica alpina C55b
                       |         Piddingtonia nummularia C55b
                       |         Queria hispanica C55b
                       |         Hohenackeria bupleurifolia C55b
                       |         Vandellia C55b
                       |           |--V. crustacea C55b
                       |           `--V. diffusa C55b
                       |         Manusiris granularis C55b
                       |         Malachra radiata C55b
                       |         Coleus africanus C55b
                       |         Micropyxis pumila [incl. Centunculus pentandrus] C55b
                       |         Elythraria fasciculata C55b
                       |         Lubinia spathulata C55b
                       |         Colea ramiflora C55b
                       |         Cirrhopetalum thouarsii C55b
                       |         Riedleia concatenata C55b
                       |         Biophytum sensitivum C55b
                       |         Stachytarpha indica C55b
                       |         Helopus annulatus [=Eriochloa annulata] C55b
                       |         Buchenavia capitata SWK87
                       |         Petitia domingensis SWK87
                       |         Tetrazygia elaeagnoides SWK87
                       |         Exothea paniculata SWK87
                       |         Matayba domingensis SWK87
                       |         Laetia procera SWK87
                       |         Torrubia fragrans SWK87
                       |         Byrsonia spicata SWK87
                       |         Triphasia trifolia SWK87
                       |         Ventarura lyonii FT05
                       |         Enterolobium F55
                       |           |--E. cyclocarpum F55
                       |           `--E. timbouva I-BH-O10
                       |         Glenniea philippinensis P88
                       |         Dryobalanops aromatica P88
                       |         Anthocephalus Pau03
                       |           |--A. cadamba Pau03
                       |           `--A. chinensis P88
                       |         Adinandra dumosa P88
                       |         Ploarium alternifolium P88
                       |         Meliopsis usambarensis RS99
                       |         Paris quadrifolia D51
                       |         Procopiana insularis S98
                       |         Gymnocarpos decander AGF98
                       |         Anabasis articulata AGF98
                       |         Dinebra retroflexa GR98
                       |         Guiera senegalensis HK98
                       |         Boscia HK98
                       |           |--B. angustifolia HK98
                       |           `--B. senegalensis HK98
                       |         Serenoa repens CS90
                       |         Plectrachne SM90
                       |           |--P. rigidissima G04
                       |           `--P. schinzii SM90
                       |         Leptosema chambersii SM90
                       |         Vetiveria elongata WF01
                       |         Tapeinia CL85
                       |           |--T. magellanica D03
                       |           `--T. pumila CL85
                       |         Rhabdotarachnoides Haupt 1957 Du07, H57
                       |           `--*R. simoni Haupt 1957 H57
                       |         Caia BC98
                       |         Albertia Schimper in Voltz 1837 FT93
                       |         Swida australis CK05
                       |         Nanophyton erinaceus TLT98
                       |         Protohyenia janovii Ananiev 1957 G97
                       |         Chlidanophyton Gensel 1973 G97
                       |         Sterulia longiflora J06
                       |         Champereia manillana J06
                       |         Faidherbia albida GC05
                       |         Aequitriradites VB02
                       |           |--A. hispidus VB02
                       |           |--A. spinulosus VB02
                       |           `--A. verrucosus VB02
                       |         Biretisporites potoniaei Delcourt & Sprumont 1955 YB02
                       |         Foraminisporis TMD02
                       |           |--F. asymmetricus VB02
                       |           |--F. tribulosus VB02
                       |           `--F. wonthaggiensis TMD02
                       |         Pilosisporites notensis TMD02
                       |         Foveosporites canalis Balme 1957 YB02
                       |         Gabonisporis YB02
                       |           |--G. cristata (Stanley) Sweet 1986 YB02
                       |           |--G. labyrinthus Srivastava 1972 YB02
                       |           `--G. vigourouxii Boltenhagen 1967 YB02
                       |         Hamulatisporis YB02
                       |           |--H. amplus Stanley 1965 YB02
                       |           `--H. hamulatis Krutzsch 1959 YB02
                       |         Intertriletes scrobiculatus Anderson 1960 YB02
                       |         Klukisporites YB02
                       |           |--K. foveolatus Pocock 1964 YB02
                       |           |--K. punctatus Venkatachala 1969 YB02
                       |           |--K. scaberis VB02
                       |           |--K. variegatus VB02
                       |           `--K. venkatachalae VB02
                       |         Lusatisporites dettmanniae (Drugg) Srivastava 1972 YB02
                       |         Osmundacidites YB02
                       |           |--O. minutus VB02
                       |           `--O. wellmanii Couper 1953 YB02
                       |         Retitriletes YB02
                       |           |--R. circolumenus VB02
                       |           `--R. nidus Srivastava 1972 YB02
                       |         Triporoletes YB02
                       |           |--T. asper Srivastava 1972 YB02
                       |           |--T. cenomanianus (Agasie) Srivastava 1977 YB02
                       |           |--T. radiatus (Dettmann) Playford 1971 YB02
                       |           |--T. reticulatus (Pocock) Playford 1971 YB02
                       |           `--T. simplex VB02
                       |         Alsophilidites bellus VB02
                       |         Concavisporites crassus VB02
                       |         Concavissimisporites VB02
                       |           |--C. apiverrucata VB02
                       |           |--C. kutchensis VB02
                       |           |--C. penolaensis VB02
                       |           |--C. variverrucosus VB02
                       |           `--C. verrucosus VB02
                       |         Contignisporites VB02
                       |           |--C. cooksoniae VB02
                       |           |--C. fornicatus VB02
                       |           |--C. glebulentus VB02
                       |           `--C. multimuratus VB02
                       |         Cooksonites variabilis VB02
                       |         Murospora florida VB02
                       |         Impardecispora VB02
                       |           |--I. apiverrucata VB02
                       |           `--I. indica VB02
                       |         Lycopodiacidites VB02
                       |           |--L. asperatus VB02
                       |           |--L. dettmanniae VB02
                       |           `--L. keupperi DV06
                       |         Foveotriletes parviretus VB02
                       |         Neoraistrickia truncata VB02
                       |         Equisetosporites VB02
                       |           |--E. concinnus VB02
                       |           |--E. eocensis VB02
                       |           |--E. virginiaensis VB02
                       |           `--E. volutus (Stanley 1965) Farabee & Canright 1986 ChrPr02
                       |         Santhalisporites imperfectus VB02
                       |         Januasporites spiniferus VB02
                       |         Schizaeoisporites eocenicus VB02
                       |         Undulatisporites pannuceus VB02
                       |         Callispora potoniei VB02
                       |         Podosporites VB02
                       |           |--P. microsaccatus VB02
                       |           `--P. tripakshi VB02
                       |         Lametatriletes indicus VB02
                       |         Ceratosporites VB02
                       |           |--C. equalis VB02
                       |           `--C. helidonensis VB02
                       |         Microcachryidites antarcticus VB02
                       |         Duplexisporites problematicus VB02
                       |         Polycingulatisporites crenulatus VB02
                       |         Baculatisporites comaumensis VB02
                       |         Matonisporis cooksoniae VB02
                       |         Stereisporites pocockii VB02
                       |         Callialasporites VB02
                       |           |--C. dampieri VB02
                       |           `--C. turbatus VB02
                       |         Boseisporites insignites VB02
                       |         Taurocusporites segmentatus VB02
                       |         Obtusisporis yarragadensis VB02
                       |         Dettmannites attenuarus VB02
                       |         Biformaesporites baculosus VB02
                       |         Triletes tuberculiformis VB02
                       |         Staplinisporites imperfectus VB02
                       |         Microreticulatisporites uniformis VB02
                       |         Coniatisporites haradensis VB02
                       |         Densoisporites mesozoicus VB02
                       |         Crassimonoletes surangei VB02
                       |         Dictyotosporites complex VB02
                       |         Appendicisporites bilateralis VB02
                       |         Carolinapollis Christopher & Prowell 2002 ChrPr02
                       |           `--*C. triangularis Christopher & Prowell 2002 ChrPr02
                       |         Holkopollenites Fairchild in Stover, Elsik & Fairchild 1966 ChrPr02
                       |           |--*H. chemardensis Fairchild in Stover, Elsik & Fairchild 1966 ChrPr02
                       |           `--H. irregularis Christopher & Prowell 2002 ChrPr02
                       |         Sparganiaceaepollenites Thiergart 1937 ChrPr02
                       |           |--*S. polygonalis Thiergart 1937 ChrPr02
                       |           |--S. magnoides Krutzsch 1970 ChrPr02
                       |           `--S. uniformis Christopher & Prowell 2002 ChrPr02
                       |         Striaperipollis Krutzsch 1966 ChrPr02
                       |           `--*S. gracilistriatus Krutzsch 1966 ChrPr02
                       |         Interpollis supplingensis (Pflug) Krutzsch 1960 ChrPr02
                       |         Plicatopollis cretacea Frederiksen & Christopher 1978 ChrPr02
                       |         Longaevipollis Khlonova 1961 ChrPr02
                       |           |--*L. sibiricus Khlonova 1961 ChrPr02
                       |           |--L. fortis Christopher & Prowell 2002 ChrPr02
                       |           `--L. subtilis Christopher & Prowell 2002 ChrPr02
                       |         Trisectoris costatus Tschudy 1970 ChrPr02
                       |         Osculapollis aequalis Tschudy 1975 ChrPr02
                       |         Libopollis jarzenii Farabee, Daghlian et al. 1984 ChrPr02
                       |         Plicapollis usitatus Tschudy 1975 ChrPr02
                       |         Monocirculopollis Krutzsch 1966 ChrPr02
                       |         Orapollis Krutzsch 1966 ChrPr02
                       |         Vaclavipollis Krutzsch 1966 ChrPr02
                       |         Vancampopollenites Kedves & Pittau 1976 ChrPr02
                       |         Gomphrenipollis Anzotegui & Cuadrado 1996 ChrPr02
                       |         Iberipollis Krutzsch 1966 ChrPr02
                       |         Papillipollis Pflug 1953 ChrPr02
                       |         Momipites Wodehouse 1933 ChrPr02
                       |           |--M. actinus Nichols & Orr 1978 ChrPr02
                       |           |--M. leffingwellii Nichols 1973 ChrPr02
                       |           |--M. rotundus (Leffingwell 1971) Nichols 1973 ChrPr02
                       |           |--M. tenuipolus Anderson 1960 ChrPr02
                       |           |--M. triorbicularis (Leffingwell 1971) Nichols 1973 ChrPr02
                       |           `--M. ventoifluminis Nichols 1973 ChrPr02
                       |         Cingutriletes clavus VB02
                       |         Coptospora VB02
                       |           |--C. microgranulosa VB02
                       |           `--C. paradoxa (Cookson & Dettmann) Dettmann 1963 RC02
                       |         Cyclosporites hughesii (Cookson & Dettmann 1958) Cookson & Dettmann 1959 RC02
                       |         Perinopollenites elatoides Couper 1958 RC02
                       |         Droseridites senonicus VH02
                       |         Triorites africaensis VH02
                       |         Gnetaceaepollenites diversus VH02
                       |         Maravenites Muller, de Di Giacomo & Van Erve 1987 ChrPr02
                       |         Reticulatisporites pudens VB02
                       |         Ruffordiaspora VB02
                       |           |--R. australiensis VB02
                       |           `--R. ludbrookiae VB02
                       |         Rugotriletes costatus (Dijkstra) Potonié 1956 TMD02
                       |         Tholisporites chulus W97
                       |         Curatella americana V-MR-M03
                       |         Aulisporites artigmosus DDL04
                       |         Camnosporites secatus DDL04
                       |         Duplicisporites DDL04
                       |           |--D. granulatus (Leschik 1956) DV06
                       |           `--D. verrucosus (Leschik 1956) DV06
                       |         Enzonalasporites DDL04
                       |           |--E. densus DDL04
                       |           |--E. ignacii DDL04
                       |           `--E. vigens Leschik 1956 DV06
                       |         Samaropollenites speciosus DDL04
                       |         Narenga porphorycoma BTT02
                       |         Cardiopteridium spitsbergense ZP86
                       |         Koeppenia Schweitzer 2000 Sc00
                       |           `--*K. eifeliensis Schweitzer 2000 Sc00
                       |         Sartilmania jabachensis Sc00
                       |         Halocarpus RL96
                       |           |--H. bidwillii RL96
                       |           `--H. biformis W91
                       |         Leonohebe mooreae W91
                       |         Pilosella officinale PAE98
                       |         Houstonia purpurea EBS98
                       |         Trichogoniopsis adenantha M-FR08
                       |         Chrysodium lanzeanum S89
                       |         Plenasium lignitum S89
                       |         Benizia calopteris S89
                       |         Graevia crenata S89
                       |         Apeibopsis gaudinii S89
                       |         Balthasaria schliebenii DL94
                       |         Gynotroches axillaris C78
                       |         Teysmanniodendron bogoriense C78
                       |         Intsia bijuga C78
                       |         Guevina avellana D03
                       |         Mitraria coccinea D03
                       |         Asteranthera chiloensis D03
                       |         Sarmienta repens D03
                       |         Decostea D03
                       |           |--D. racemosa D03
                       |           `--D. ruscifolia D03
                       |         Rhaphithamnus cyanocarpus D03
                       |         Cynoctonum pachyphyllum D03
                       |         Myoschilos oblongus D03
                       |         Schoenodon chilensis D03
                       |         Saxegothea conspicua D03
                       |         Susarium segethi D03
                       |         Hebenstretia angolensis B06
                       |         Laurofillum JD05
                       |           |--L. acutimontanum JD05
                       |           `--L. pseudoprinceps JD05
                       |         Viscophyllum weylandii JD05
                       |         Icaciniphyllum artocarpites JD05
                       |         Hamatophyton verticillatum CWC87
                       |         Lepidostrobophyllum fimbriatum CWC87
                       |         Neocystites Barrande 1887 [=Neocystis Bather 1889] K67
                       |         Piptadenia buchananii W86
                       |         Hopoea odorata B88
                       |         Symphiocarpos B75
                       |           |--S. occidentalis B75
                       |           `--S. orbiculatus B75
                       |         Drepanostachyum intermedium Da07
                       |         Maotia puya Da07
                       |         Alcimandra cathcartii Da07
                       |         Bonnamia latifolia Par03
                       |         Gmelina arborea Pau03
                       |         Loricaria columbiana R96
                       |         Hesperomeles ferruginea R96
                       |         Neurolepis arista R96
                       |         Amphipterygium adstringens G69
                       |         Calycotome RBA00
                       |           |--C. spinosa C74
                       |           `--C. villosa RBA00
                       |         Mariopteris pygmea R79
                       |         Lagarostrobos SN08
                       |           |--L. colensoi RL96
                       |           `--L. franklanii SN08
                       |         Befaria racemosa M83
                       |         Proserpinaca palustris M83
                       |         Hepatica transsilvanica H09
                       |         Pituranthus triradiatus HF03
                       |         Dubautia linearis MK08
                       |         Fagaropsis angolensis K09
                       |         Stricklandia Buckman 1845 AB65
                       |         Bedfordia salicina SN08
                       |         Megalodonta beckii B97
                       |         Crysophila guagara BM76
                       |         Paranymphaea crassifola OB13
                       |         Stranvaesia niitakayamensis ChuPe02
                       |         Guazuma tomentosa B14
                       |         Thyrsopteris [Thyrsopteridaceae] P-GR14
                       |           `--T. elegans P-GR14
                       |         Pringlea antiscorbutica SBG97
                       |         Petalostylis labicheoides ML13
                       |         Argusia argentea NC91
                       |         Moerua rigida CS77
                       |         Bolboschoenus maritimus CS77
                       |         Antenostoma fasciculatum BT72
                       |         Brickiella californica BT72
                       |         Parmentiera alata BTA75
                       |         Bouchea prismatica BTA75
                       |         Diplopeltis stuartii MLP09
                       |         Carrichtera annua K10
                       |         Hammada L98
                       |           |--H. salicornica L98
                       |           `--H. scoparia L98
                       |         Anvillea garcini L98
                       |         Rhacophyllum coriaceum R87
                       |         Euodia M87
                       |           |--E. haplophylla WO95
                       |           `--E. micrococca M87
                       |         Noeggerathiopsis S87
                       |           |--N. media S87
                       |           |--N. prisca S87
                       |           `--N. spathulata S87
                       |         Fortunella japonica B00
                       |         Acradenia euodiiformis B00
                       |         Armeniaca RJ11
                       |           |--A. sibirica RJ11
                       |           `--A. vulgaris TW07
                       |         Peritoma serrulatum R35
                       |         Camphora officinalis C70
                       |         Gonioma kamassi P65
                       |         Orthodicranum montanum N77
                       |         Lepidophyllum quadrangulare RJ11
                       |         Thalia geniculata F11
                       |         Pullea stutzeri WO95
                       |         Coleospermum paniculatum WO95
                       |         Galax aphylla R56
                       |         Cheilotetras caledonica Wellman & Richardson 1993 SP12
                       |         Tetrahedraletes Strother & Traverse 1979 SP12
                       |           `--T. medinensis (Strother & Traverse) Wellman & Richardson 1993 SP12
                       |         Rimosotetras problematica Burgess 1991 SP12
                       |         Imperfectotriletes Steemans et al. 2000 SP12
                       |           |--I. patinatus Steemans et al. 2000 SP12
                       |           `--I. varvdovae (Richardson) Steemans et al. 2000 SP12
                       |         Pseudodyadospora laevigata Johnson 1985 SP12
                       |         Gneudnaspora divellomedia (Chibrikova) Balme 1988 [=Laevolancis divellomedia] SP12
                       |           |--G. d. var. divellomedia SP12
                       |           `--G. d. var. minor Breuer et al. 2007 [=Laevolancis divellomedia var. minor] SP12
                       |         Laevolancis chibrikovae Steemans et al. 2000 [=Gneudnaspora chibrikovae] SP12
                       |         Dyadospora Strother & Traverse 1979 SP12
                       |           |--D. murusattenuata Strother & Traverse 1979 SP12
                       |           `--D. murusdensa Strother & Traverse 1979 SP12
                       |         Abditusdyadus Wellman & Richardson 1996 SP12
                       |           `--A. laevigatus Wellman & Richardson 1996 SP12
                       |         Acontotetras inconspicuis Richardson 1996 SP12
                       |         Ancyrospora ancyrea Richardson 1964 SP12
                       |         Archaeozonotriletes chulusnanus Steemans et al. 1996 SP12
                       |         Artemopyra Burgess & Richardson 1991 SP12
                       |           `--A. brevicosta Burgess & Richardson 1991 SP12
                       |         Breconisporites Richardson et al. 1982 SP12
                       |         Chelinohilates Richardson 1996 SP12
                       |           `--C. erraticus Richardson 1996 SP12
                       |         Chelinospora Allen 1965 SP12
                       |           |--C. cantabrica Richardson et al. 2001 SP12
                       |           |--C. cassicula Richardson & Lister 1969 SP12
                       |           `--C. poecilomorpha (Richardson & Ioannides) Richardson et al. 2001 SP12
                       |         Clivosispora verrucata McGregor & Camfield 1976 SP12
                       |           |--C. v. var. verrucata SP12
                       |           `--C. v. var. convoluta McGregor & Camfield 1976 SP12
                       |         Coronaspora Rodríguez 1978 SP12
                       |           `--C. cromatica Jansonius & Hills 1979 SP12
                       |         Cymbohilates Richardson 1996 SP12
                       |           |--C. baqaensis Breuer et al. 2007 SP12
                       |           |--C. cymosus Richardson 1996 SP12
                       |           `--C. disponerus Richardson 1996 SP12
                       |         Cymbosporites Allen 1965 SP12
                       |           |--C. dammamensis Steemans 1995 SP12
                       |           |--C. proteus McGregor & Camfield 1976 SP12
                       |           `--C. stellospinosus Steemans 1989 SP12
                       |         Dictyotriletes Naumova ex Ischchenko 1952 SP12
                       |           `--D. subgranifer McGregor 1973 SP12
                       |         Hispanaediscus Cramer 1966 SP12
                       |           `--H. verrucatus (Cramer) Burgess & Richardson 1991 SP12
                       |         Hystricosporites McGregor 1960 SP12
                       |         Rugosisporites Dufka 1995 SP12
                       |         Scylaspora Burgess & Richardson 1995 SP12
                       |         Segestrespora Burgess 1991 SP12
                       |           |--S. laevigata Burgess 1991 SP12
                       |           `--S. membranifera (Johnson) Burgess 1991 SP12
                       |         Stegambiquadrella contenta Johnson 1985 SP12
                       |         Velatitetras Burgess 1991 SP12
                       |           |--V. laevigata Burgess 1991 SP12
                       |           `--V. retimembrana (Miller & Eames 1982) Wellman & Richardson 1996 SP12
                       |         Zonotriletes Luber & Walts 1938 SP12
                       |         Bathurstia Hueber 1972 SP12
                       |         Estinnophyton Fairon-Demaret 1978 SP12
                       |         Hollandophyton Rogerson et al. 2002 SP12
                       |         Junggaria Dou & Sun 1983 SP12
                       |         Metacladophyton Wang & Geng 1997 SP12
                       |         Cirratriradites diaphanus Steemans 1989 SP12
                       |         Sphaerasaccus glabellus Steemans et al. 2000 SP12
                       |         Amicosporites streelii Steemans 1989 SP12
                       |         Brochotriletes foveolatus Naumova 1953 SP12
                       |         Latosporites ovalis Breuer et al. 2007 SP12
                       |         Camarozonotriletes sextantii McGregor & Camfield 1976 SP12
                       |         Samarisporites triangulatus Allen 1965 SP12
                       |         Lophozonotriletes media Taugourdeau-Lantz 1967 SP12
                       |         Cyatheacidites annulatus V12
                       |         Cibotiidites tuberculiformis V12
                       |         Peromonolites bowenii V12
                       |         Dacrydiumites praecupressinoides V12
                       |         Dodartia orientalis Si00
                       |         Echinopanax elatum Si00
                       |         Evodia roxburghiana Si00
                       |         Turraeanthus africana Si00
                       |         Doxantha unguis-cati BT87
                       |         Atamisquea emarginata BT87
                       |         Trixus californicus BT87
                       |         Viquiera stenoloba BT87
                       |         Pleioblastus chino TM96
                       |--Bryata MS98
                       `--Pan-Tracheophyta [Polysporangiomorpha] CD07
                            |--Horneophyton lignieri D98, BC98
                            `--Apo-Tracheophyta WM14

Streptophyta incertae sedis:
  Sycidiaceae E03
  Umbellinaceae E03
  Columbella ningbingensis E03
  *Trochiliscus panderi [=Miliola panderi Ehrenberg 1859] E03
  Porocharaceae WH78
  Clavator [Clavatoraceae] WH78
    `--C. reidi Harris 1939 WH78
  Praechara Birina 1948 WH78
  Raskyella sahariana Mebrouk 1993 GH99
  Neochara huananensi NG13
  Grovesichara changzhouensis NG13
  Obtusochara prisca NG13

Nomina nuda: Chrysodium doriae Squinabol 1889 S89
             Nassauvia dusenii Hoffm. in Dusén 2003 D03

Chaetosphaeridium [Chaetosphaeridiobiotes, Chaetosphaeridiophyceae, Chaetosphaeridiophyta] KM01

Streptophyta [Charophyceae, Charophyta, Eophyceae, Phragmophytina, Rudophytae, Stichophycidae, Streptobionta, Zygnemobiotes] WM14

*Type species of generic name indicated

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