Reconstruction of Horneophyton lignieri, copyright Falconaumanni.

Belongs within: Streptophyta.
Contains: Cingulati, Zonati, Pollenites, Apo-Tracheophyta, Marchantiophyta, Bryophyta, Anthocerotopsida.

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. Sperm cells are differentiated from eggs, and some species will have sperm and eggs produced by distinct individuals. Ciliated basal bodies, when present, have a distinctive multilayered structure of microtubules and cytoskeletal anchor. Adjacent vegetative cells are connected by junctions (plasmodesmata) perforating the cell wall (Adl et al. 2019).

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. Non-vascular plants include the mosses, liverworts and hornworts (Anthocerotophyta), with the hornworts being small non-vascular plants that produce relatively long horn-like sporophytes. Hornwort gametophytes are thalloid and often contain symbiotic Nostoc cyanobacteria. Leiosporoceros dussii represents the sister taxon to other living hornworts, with distinctive features including the production of monolete spores exclusively in isobilateral tetrads, large, irregularly thickened pseudoelaters embedded in a mucilaginous matrix, and occurrence of Nostoc symbionts in longitudinally aligned, branching strands rather than spherical colonies (Stotler & Crandall-Stotler 2005). This species is known from Central America and the Antilles.

There has been much debate over the exact interrelationships between the vascular and non-vascular plants. Many authors place the hornworts 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, whereas 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. The earliest Ordovician spores, referred to as cryptospores, are most commonly associated in dyads and tetrads and characterised by the lack of trilete marks. The ornament on the outer spore wall or membrane is inconspicuous in the earliest exemplars but tends to become more developed over time. Trilete spores first appear in the later Ordovician (Steemans et al. 2012). 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 earlier 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. Vegetative propagation is also possible in many species. 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 lignieri 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).

Aequitriradites: the mark of the Cretaceous
Published 2 September 2014
Diagram of Aequitriradites ornatus, from Upshaw (1963).

The Cretaceous period is best known in popular culture as the time of Tyrannosaurus and Triceratops, of Pteranodon and Quetzalcoatlus, of Elasmosaurus and mosasaurs. But it was also, in an arguably even more significant way, the time of Aequitriradites.

Aequitriradites is the fossil represented in the diagram at the top of this post. It is not very large: at its largest, the species shown above is about a tenth of a millimetre across (Upshaw 1963). It is, in fact, the spore from a liverwort. The vegetative parts of liverworts mostly do not have much of a fossil record, being soft and prone to decay, though long-time readers may recall a suggestion that this spotty fossil record was occasionally dramatic (alas, general opinion seems to have not been swayed). However, their spores are more resistent, and hence may be abundant as fossils. Because it is rarely possible to tell exactly which plant they came from, fossil spores (and pollen) are classified as form taxa, parallel to the classification of other plant fossils. Aequitriradites species are characterised by a membranous flange (a zona) running around the outside of the spore, together with a triradiate laesura pattern (the fissures that mark where the spore opens when it germinates) on one face. Depending on the species, the laesurae may be well-marked or faint. There may also be an opening in the spore wall at the apex of the face opposite the laesurae (Cookson & Dettmann 1961). It has been suggested that the otherwise unidentified liverworts that produced Aequitriradites spores were probably related to the modern liverwort order Sphaerocarpales, and Archangelsky & Archangelsky (2005) compared Aequitriradites to the spores of the aquatic genus Riella.

Alternate faces of specimens of Aequitriradites plicatus, from Archangelsky & Archangelsky (2005).

The abundance of plant spores and pollen is such that they are commonly used as ‘index fossils’, indicators of the age of the rock they are found in. Aequitriradites contains various species throughout the Cretaceous period (species assigned to this genus from the Triassic seem to have since been re-classified). Li (2014) identified the appearance of Aequitriradites spinulosus in the very latest Jurassic as one of the better indicators of the start of the Cretaceous period in the Qinghai-Xizang Plateau in China. Aequitriradites species seem to have been most abundant in the Early Cretaceous, becoming rarer in the Late Cretaceous. Establishing the latest appearance of a spore taxon in the fossil record can be difficult, because of the possibility of re-working (fossil spores being disassociated from their original deposit and re-buried in a later one), but non-reworked examples of Aequitriradites in the latter part of the Late Cretaceous were alluded to by Askin (1990). TLDR: If you’ve got Aequitriradites, you’ve got Cretaceous.

Systematics of Embryophyta
<==Embryophyta [Acotyledones, Cormophyta, Cryptogamae, Embryobiotes, Hepatophyta]WM14
|--Pan-Tracheophyta [Polysporangiomorpha]CD07
| |--Apo-TracheophytaWM14
| `--Horneophyton [Horneophytaceae, Horneophytales, Horneophytopsida]D98
| `--H. lignieri (Kidston & Lang) Barghoorn & Darrah 1938C93
| i. s.: Cryptocoleopsis imbricataB-OO10
| Hydrocryphaea wardiiB-OO10
| Prasanthus suecicusB-OO10
| Yunnanobryon rhyacophilumB-OO10
| Lobatiriccardia yunnannesisB-OO10
| Gottschelia grolleiB-OO10
| Hamatostrepta concinnaB-OO10
| `--BryophytaWM14
`--Anthocerotophyta [Anthocerotae]WM14
`--Leiosporoceros Hässel 1986 [Leiosporocerotaceae, Leiosporocerotales, Leiosporocerotopsida]SC-S05
`--L. dussiiSC-S05

Embryophyta incertae sedis:
Eophyllophyton bellumK01
|--S. mirabileZA02
`--S. wusihense [=Lepidodendron wusihense, Eolepidodendron wusihense]CWC87
Colophospermum mopaneWS02
|--I. abundans Tosolini & McLoughlin 2002TMD02
`--I. daharensis Barale 1999BO02
Ptilophyllum caucasicumG10
Eranthis hyemalisV02
Synsepalum dulcificumZB03
Rhaciopteris busseanaA38
Diplothmema subfurcatumA38
Megalopteris dawsoniA38
Sporangites acuminataA38
Pterispermostrobus bifurcatusA38
|--W. concinnaA38
`--W. dawsonianaA38
|--C. baileyiA38
|--C. cornutumA38
|--C. crampiA38
`--C. obliquumA38
|--A. acicularisA38
|--A. equisetiformisG31
`--A. parvulus [incl. A. grandis]A38
Campsiandra laurifoliaDS86
Radiola linoides [=Linum radiola]C55a
Succowia balearica [=Bunias balearica]C55a
|--D. bulbiferaC55a
|--D. diphyllaJ83
|--D. enneaphyllaC55a
`--D. laciniataJ83
Aretia villosaC55a
Nauenburgia thyrsifloraC55a
Cybistax antisyphiliticaC55a
Nonnea ventricosaC55a
Doratanthera linearisC55a
Ramphicarpa fistulosaC55a
Anodendron paniculatumC55a
Lachnopylis oppositifoliaC55a
Hugueninia tanacetifoliaC55a
Adenostyles leucophyllaC55a
Kernera saxatilisC55a
Petrocallis pyrenaicaC55a
Orobus luteusC55a
Serratula nudicaulisC55a
Betonica alopecurusC55a
Zephyritis taitensisC55a
Tristegis glutinosaC55b
Cucubalus bacciferusC55b
|--C. aromaticumC55b
`--C. aureumC55b
Hyssopus officinalisC55b
|--B. polymorphumC55b
`--B. virgatumC55b
Anacharis alsinastrumC55b
|--E. hirsutumC55b
`--E. lensC55b
Lappa majorC55b
|--A. ambrosioidesD03
`--A. botrysC55b
Agathophytum bonus-henricus [=Blitum bonus-henricus]C55b
Guilandina bonduc [incl. G. bonducella]C55b
Drepanocarpus lunatusC55b
Ecastaphyllum browneiC55b
Schwenkia americanaC55b
|--T. brasilianaC55b
|--T. frutescensC55b
`--T. maritimaC55b
Ptarmica alpinaC55b
Piddingtonia nummulariaC55b
Queria hispanicaC55b
Vandellia crustaceaC55b
Manusiris granularisC55b
|--M. capitataA73
`--M. radiataC55b
Micropyxis pumila [incl. Centunculus pentandrus]C55b
Elythraria fasciculataC55b
Lubinia spathulataC55b
Colea ramiflora DC. 1845 (see below for synonymy)S77
Riedleia concatenataC55b
Biophytum sensitivumC55b
|--S. indicaC55b
`--S. jamaicensisFS90
Buchenavia capitataSWK87
Exothea paniculataSWK87
Matayba domingensisSWK87
Laetia proceraSWK87
Torrubia fragransSWK87
Byrsonia spicataSWK87
Triphasia trifoliaSWK87
Ventarura lyoniiFT05
Glenniea philippinensisP88
|--A. cadambaPau03
`--A. chinensisP88
Ploarium alternifoliumP88
Meliopsis usambarensisRS99
Paris quadrifoliaD51
Procopiana insularisS98
Gymnocarpos decanderAGF98
Dinebra retroflexaGR98
Leptosema chambersiiSM90
|--T. magellanicaD03
`--T. pumilaCL85
Rhabdotarachnoides Haupt 1957Du07, H57
`--*R. simoni Haupt 1957H57
Albertia Schimper in Voltz 1837FT93
Swida australisCK05
Nanophyton erinaceusTLT98
Chlidanophyton Gensel 1973G97
Sterulia longifloraJ06
Champereia manillanaJ06
|--A. hispidusVB02
|--A. spinulosusVB02
`--A. verrucosusVB02
Biretisporites potoniaei Delcourt & Sprumont 1955YB02
|--F. asymmetricusVB02
|--F. tribulosusVB02
`--F. wonthaggiensisTMD02
Pilosisporites notensisTMD02
|--F. canalis Balme 1957YB02
`--F. insculptusP63
|--G. cristata (Stanley) Sweet 1986YB02
|--G. labyrinthus Srivastava 1972YB02
`--G. vigourouxii Boltenhagen 1967YB02
|--H. amplus Stanley 1965YB02
`--H. hamulatis Krutzsch 1959YB02
Intertriletes scrobiculatus Anderson 1960YB02
|--K. foveolatus Pocock 1964YB02
|--K. punctatus Venkatachala 1969YB02
|--K. scaberisVB02
|--K. variegatusVB02
`--K. venkatachalaeVB02
Lusatisporites dettmanniae (Drugg) Srivastava 1972YB02
|--O. minutusVB02
`--O. wellmanii Couper 1953YB02
|--R. circolumenusVB02
`--R. nidus Srivastava 1972YB02
|--T. asper Srivastava 1972YB02
|--T. cenomanianus (Agasie) Srivastava 1977YB02
|--T. radiatus (Dettmann) Playford 1971YB02
|--T. reticulatus (Pocock) Playford 1971YB02
`--T. simplexVB02
Alsophilidites bellusVB02
Concavisporites crassusVB02
|--C. apiverrucataVB02
|--C. kutchensisVB02
|--C. penolaensisVB02
|--C. variverrucosusVB02
`--C. verrucosusVB02
|--C. cooksoniaeVB02
|--C. fornicatusVB02
|--C. glebulentusVB02
`--C. multimuratusVB02
Cooksonites variabilisVB02
|--M. auritaP63
|--M. conduplicataP63
|--M. dubiaP63
|--M. floridaVB02
|--M. friendiiP63
|--M. intortaP63
|--M. strigataP63
|--M. sublobataP63
`--M. tripulvinata Staplin 1960P63
|--I. apiverrucataVB02
`--I. indicaVB02
|--L. asperatusVB02
|--L. dettmanniaeVB02
`--L. keupperiDV06
Foveotriletes parviretusVB02
Neoraistrickia truncataVB02
|--E. concinnusVB02
|--E. eocensisVB02
|--E. virginiaensisVB02
`--E. volutus (Stanley 1965) Farabee & Canright 1986ChrPr02
Santhalisporites imperfectusVB02
Januasporites spiniferusVB02
Schizaeoisporites eocenicusVB02
Undulatisporites pannuceusVB02
Callispora potonieiVB02
|--P. microsaccatusVB02
`--P. tripakshiVB02
Lametatriletes indicusVB02
|--C. equalisVB02
`--C. helidonensisVB02
Microcachryidites antarcticusVB02
Duplexisporites problematicusVB02
Polycingulatisporites crenulatusVB02
Baculatisporites comaumensisVB02
Matonisporis cooksoniaeVB02
Stereisporites pocockiiVB02
|--C. dampieriVB02
`--C. turbatusVB02
Boseisporites insignitesVB02
Taurocusporites segmentatusVB02
Obtusisporis yarragadensisVB02
Dettmannites attenuarusVB02
Biformaesporites baculosusVB02
Triletes tuberculiformisVB02
Staplinisporites imperfectusVB02
|--M. lunatusP63
`--M. uniformisVB02
Coniatisporites haradensisVB02
Densoisporites mesozoicusVB02
Crassimonoletes surangeiVB02
Dictyotosporites complexVB02
Appendicisporites bilateralisVB02
Carolinapollis Christopher & Prowell 2002ChrPr02
`--*C. triangularis Christopher & Prowell 2002ChrPr02
Holkopollenites Fairchild in Stover, Elsik & Fairchild 1966ChrPr02
|--*H. chemardensis Fairchild in Stover, Elsik & Fairchild 1966ChrPr02
`--H. irregularis Christopher & Prowell 2002ChrPr02
Striaperipollis Krutzsch 1966ChrPr02
`--*S. gracilistriatus Krutzsch 1966ChrPr02
Interpollis supplingensis (Pflug) Krutzsch 1960ChrPr02
Plicatopollis cretacea Frederiksen & Christopher 1978ChrPr02
Longaevipollis Khlonova 1961ChrPr02
|--*L. sibiricus Khlonova 1961ChrPr02
|--L. fortis Christopher & Prowell 2002ChrPr02
`--L. subtilis Christopher & Prowell 2002ChrPr02
Trisectoris costatus Tschudy 1970ChrPr02
Osculapollis aequalis Tschudy 1975ChrPr02
Libopollis jarzenii Farabee, Daghlian et al. 1984ChrPr02
Monocirculopollis Krutzsch 1966ChrPr02
Orapollis Krutzsch 1966ChrPr02
Vaclavipollis Krutzsch 1966ChrPr02
Vancampopollenites Kedves & Pittau 1976ChrPr02
Gomphrenipollis Anzotegui & Cuadrado 1996ChrPr02
Iberipollis Krutzsch 1966ChrPr02
Papillipollis Pflug 1953ChrPr02
Cingutriletes clavusVB02
|--C. microgranulosaVB02
`--C. paradoxa (Cookson & Dettmann) Dettmann 1963RC02
Cyclosporites hughesii (Cookson & Dettmann 1958) Cookson & Dettmann 1959RC02
Perinopollenites elatoides Couper 1958RC02
Droseridites senonicusVH02
Gnetaceaepollenites diversusVH02
Maravenites Muller, de Di Giacomo & Van Erve 1987ChrPr02
|--R. cancellatusP63
|--R. peltatusP63
|--R. planus Hughes & Playford 1961P63
|--R. pudensVB02
|--R. rudis Staplin 1960P63
`--R. variolatusP63
|--R. australiensisVB02
`--R. ludbrookiaeVB02
Rugotriletes costatus (Dijkstra) Potonié 1956TMD02
Tholisporites Butterworth & Williams 1958 [Patinati]P63
|--*T. scoticus Butterworth & Williams 1958P63
|--T. chulusW97
`--T. foveolatus Hughes & Playford 1961P63
Aulisporites artigmosusDDL04
Camnosporites secatusDDL04
|--D. granulatus (Leschik 1956)DV06
`--D. verrucosus (Leschik 1956)DV06
|--E. densusDDL04
|--E. ignaciiDDL04
`--E. vigens Leschik 1956DV06
Samaropollenites speciosusDDL04
Narenga porphorycomaBTT02
Cardiopteridium spitsbergenseZP86
Koeppenia Schweitzer 2000Sc00
`--*K. eifeliensis Schweitzer 2000Sc00
Sartilmania jabachensisSc00
|--H. bidwilliiRL96
`--H. biformisW91
Leonohebe mooreaeW91
Pilosella officinalePAE98
|--H. ceruleaJ23
`--H. purpureaEBS98
Trichogoniopsis adenanthaM-FR08
Chrysodium lanzeanumSq89
Plenasium lignitumSq89
Benizia calopterisSq89
Graevia crenataSq89
Apeibopsis gaudiniiSq89
Balthasaria schliebeniiDL94
Gynotroches axillarisC78
Teysmanniodendron bogorienseC78
Intsia bijugaC78
Guevina avellanaD03
Mitraria coccineaD03
Asteranthera chiloensisD03
Sarmienta repensD03
|--D. racemosaD03
`--D. ruscifoliaD03
Rhaphithamnus cyanocarpusD03
Myoschilos oblongusD03
Schoenodon chilensisD03
Susarium segethiD03
Hebenstretia angolensisB06
|--L. acutimontanumJD05
`--L. pseudoprincepsJD05
Viscophyllum weylandiiJD05
Icaciniphyllum artocarpitesJD05
Hamatophyton verticillatumCWC87
Lepidostrobophyllum fimbriatumCWC87
Neocystites Barrande 1887 [=Neocystis Bather 1889]K67
Hopoea odorataB88
|--S. occidentalisB75
`--S. orbiculatusB75
Drepanostachyum intermediumDa07
Maotia puyaDa07
Alcimandra cathcartiiDa07
Bonnamia latifoliaPar03
Loricaria columbianaR96
Hesperomeles ferrugineaR96
Neurolepis aristaR96
Amphipterygium adstringensG69
|--C. spinosaC74
`--C. villosaRBA00
|--M. muricataG31
`--M. pygmeaR79
|--L. colensoiRL96
`--L. franklaniiSN08
Befaria racemosaM83
Pituranthus triradiatusHF03
Dubautia linearisMK08
Fagaropsis angolensisE09
Stricklandia Buckman 1845AB65
Megalodonta beckiiB97
Crysophila guagaraBM76
Paranymphaea crassifolaOB13
Stranvaesia niitakayamensisChuPe02
Guazuma tomentosaB14
Thyrsopteris [Thyrsopteridaceae]P-GR14
`--T. elegansP-GR14
Pringlea antiscorbuticaSBG97
Antenostoma fasciculatumBT72
Brickiella californicaBT72
|--H. salicornicaL98
`--H. scopariaL98
Rhacophyllum coriaceumR87
|--N. aequalisG31
|--N. mediaS87
|--N. priscaS87
`--N. spathulataS87
|--A. sibiricaRJ11
`--A. vulgarisTW07
Peritoma serrulatumR35
Camphora officinalisC70
Gonioma kamassiP65
|--L. lanceolatumG31
`--L. quadrangulareRJ11
Thalia geniculataF11
Pullea stutzeriWO95
Coleospermum paniculatumWO95
Galax aphyllaR56
Cheilotetras caledonica Wellman & Richardson 1993SP12
Tetrahedraletes Strother & Traverse 1979SP12
`--T. medinensis (Strother & Traverse) Wellman & Richardson 1993SP12
Rimosotetras problematica Burgess 1991SP12
Imperfectotriletes Steemans et al. 2000SP12
|--I. patinatus Steemans et al. 2000SP12
`--I. varvdovae (Richardson) Steemans et al. 2000SP12
Pseudodyadospora laevigata Johnson 1985SP12
Gneudnaspora divellomedia (Chibrikova) Balme 1988 [=Laevolancis divellomedia]SP12
|--G. d. var. divellomediaSP12
`--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 1979SP12
|--D. murusattenuata Strother & Traverse 1979SP12
`--D. murusdensa Strother & Traverse 1979SP12
Abditusdyadus Wellman & Richardson 1996SP12
`--A. laevigatus Wellman & Richardson 1996SP12
Acontotetras inconspicuis Richardson 1996SP12
Ancyrospora ancyrea Richardson 1964SP12
Archaeozonotriletes Naumova 1953P63
|--A. chulusLT80
`--A. chulusnanus Steemans et al. 1996SP12
Artemopyra Burgess & Richardson 1991SP12
`--A. brevicosta Burgess & Richardson 1991SP12
Breconisporites Richardson et al. 1982SP12
Chelinohilates Richardson 1996SP12
`--C. erraticus Richardson 1996SP12
Chelinospora Allen 1965SP12
|--C. cantabrica Richardson et al. 2001SP12
|--C. cassicula Richardson & Lister 1969SP12
`--C. poecilomorpha (Richardson & Ioannides) Richardson et al. 2001SP12
|--C. torquata McGregor 1973LT80
`--C. verrucata McGregor & Camfield 1976SP12
|--C. v. var. verrucataSP12
`--C. v. var. convoluta McGregor & Camfield 1976SP12
Coronaspora Rodríguez 1978SP12
`--C. cromatica Jansonius & Hills 1979SP12
Cymbohilates Richardson 1996SP12
|--C. baqaensis Breuer et al. 2007SP12
|--C. cymosus Richardson 1996SP12
`--C. disponerus Richardson 1996SP12
Cymbosporites Allen 1965SP12
|--C. dammamensis Steemans 1995SP12
|--C. proteus McGregor & Camfield 1976SP12
`--C. stellospinosus Steemans 1989SP12
Dictyotriletes Naumova ex Ischchenko 1952SP12
|--D. caperatusP63
`--D. subgranifer McGregor 1973SP12
Hispanaediscus Cramer 1966SP12
`--H. verrucatus (Cramer) Burgess & Richardson 1991SP12
Hystricosporites McGregor 1960SP12
Rugosisporites Dufka 1995SP12
Scylaspora Burgess & Richardson 1995SP12
Segestrespora Burgess 1991SP12
|--S. laevigata Burgess 1991SP12
`--S. membranifera (Johnson) Burgess 1991SP12
Stegambiquadrella contenta Johnson 1985SP12
Velatitetras Burgess 1991SP12
|--V. laevigata Burgess 1991SP12
`--V. retimembrana (Miller & Eames 1982) Wellman & Richardson 1996SP12
Zonotriletes Luber & Walts 1938SP12
|--Z. ciliato-marginatus Waltz in Luber & Waltz 1941P63
|--Z. latizonalis Waltz in Luber & Waltz 1941P63
`--Z. sulcatus Waltz in Luber & Waltz 1938P63
Bathurstia Hueber 1972SP12
Estinnophyton Fairon-Demaret 1978SP12
Hollandophyton Rogerson et al. 2002SP12
Junggaria Dou & Sun 1983SP12
Metacladophyton Wang & Geng 1997SP12
Sphaerasaccus glabellus Steemans et al. 2000SP12
Amicosporites streelii Steemans 1989SP12
Brochotriletes foveolatus Naumova 1953SP12
Latosporites ovalis Breuer et al. 2007SP12
Camarozonotriletes sextantii McGregor & Camfield 1976SP12
Samarisporites triangulatus Allen 1965SP12
Lophozonotriletes (Naumova) Potonié 1958P63
|--*L. lebedianensis Naumova 1953P63
|--L. appendices (Hacquebard & Barss) Playford 1963 [=Cincturasporites appendices Hacquebard & Barss 1957]P63
|--L. dentatus Hughes & Playford 1961P63
|--L. media Taugourdeau-Lantz 1967SP12
|--L. rarituberculatus (Luber) Kedo 1957 (see below for synonymy)P63
`--L. variverrucatus Playford 1963P63
Cyatheacidites annulatusV12
Cibotiidites tuberculiformisV12
Peromonolites boweniiV12
Dodartia orientalisSi00
|--E. elatumSi00
`--E. horridumC52
Turraeanthus africanaSi00
Atamisquea emarginataBT87
Trixus californicusBT87
Viquiera stenolobaBT87
Pleioblastus chinoTM96
Acanthyllis tragacanthoidesB28
Amberboa lippiiB28
Koniga maritimaB28
Triticosecale rimpaniPS96
Apollonias barbujanaBBO02
Eubotryoides grayanaBBO02
Acalyptospora Desm. 1848 (n. d.)KC01
Rhodocolea racemosa [incl. Bignonia compressa Lank 1785, Colea decora DC. 1845]S77
|--R. r. var. racemosaS77
`--R. r. var. humblotiana [incl. Colea aberrans Baill. 1889]S77
Dichrotrichum ternateum [=Bignonia ternatea Reinw. ex De Vriese 1858 (n. n.)]S77
Thaumatococcus danielliiW92
Discoreophyllum cumminsiiW92
Pelucha trifidaMF68
Corsinipollenites thiergartiiWPL96
Anapiculatisporites Potonié & Kremp 1954P63
|--A. concinnusP63
|--A. largusWT04
`--A. serratus Playford 1963P63
|--T. complanatus Staplin 1960P63
|--T. incisotrilobus (Naumova) Potonié & Kremp 1956P63
`--T. vetustusPSC99
Rotaspora fractaPSC99
Raistrickia nigraPSC99
|--T. batillatus Hughes & Playford 1961P63
|--T. marginatusPSC99
`--T. trivalvis (Waltz) Potonié & Kremp 1956P63
Schulzospora campylopteraPSC99
|--L. lenaensisG10
`--L. prynardiiG10
Chaja ivorensisK68
Cupuliferoidaepollenites liblarensisP92
Ulmipollenites undulosusP92
Palaeostachya arborescensG31
Pinnularia columnarisG31
Semaropsis fluitansG31
Stachannularia tuberculataG31
Collipseridium dawsonianumG31
Cardiocarpus boulayiG31
Lonchopteris alataG31
|--C. koraiensisG31
`--C. pteridiumG31
|--C. confertaG31
|--C. crassinervisG31
`--C. murenensisG31
Mixoneura subcrenataG31
Lagenospermum oblongumG31
Annulariopsis inopinataG31
Chiropteris reniformisG31
Neurogangamopteris cardiopteroidesG31
Caenodendron primaevumG31
Equisetites mirabilisG31
Mesopitys tschichatschewiG31
Psygmophyllum mongolicumG31
Glottophyllum cuneatumG31
Canthiumidites bellusOW01
Monotocidites galeatusOW01
Wittania somniferaA73
|--L. curiosusP63
|--L. inermisP63
|--L. microgranulatusP63
|--L. ornatusP63
|--L. pagius Allen 1965LT80
`--L. subintortusP63
|--L. s. var. subintortusP63
`--L. s. var. rotundatusP63
|--A. arenorugosa McGregor 1973LT80
|--A. brandtii Streel 1964LT80
|--A. perpusillus (Naum.) McGregor 1973LT80
`--A. plicata (Allen) Streel 1967LT80
Peplis [=Chabraea Adans. 1763 (nom. illeg.)]BGS05
Hofmeisteria filifoliaM69
Baeriopsis guadalupensisM69
|--A. apludoidesJ23
`--A. oligostachyumJ23
Anulatisporites Loose ex Potonié & Kremp 1954P63
|--A. anulatus (Loose) Potonié & Kremp 1954P63
|--A. canaliculatus Playford 1963P63
|--A. labiatus Hughes & Playford 1961P63
`--A. orbiculatus (Waltz) Playford 1962 (see below for synonymy)P63
Hymenozonotriletes Naumova 1937P63
|--H. auranthiacus [=Zonotriletes auranthiacus (Naumova) Waltz in Luber & Waltz 1938]P63
|--H. concavus Ishchenko 1956P63
`--H. praetervisus Naumova 1953P63
Labiadensites Hacquebard & Barss 1957P63
|--*L. attenuatus Hacquebard & Barss 1957P63
`--L. fimbriatus (Waltz) Hacquebard & Barss 1957 (see below for synonymy)P63
Spinozonotriletes Hacquebard 1957P63
|--*S. uncatus Hacquebard 1957P63
|--S. balteatus Playford 1963P63
`--S. exiguus Staplin 1960P63
Cincturasporites Hacquebard & Barss 1957P63
Euryzonotriletes Naumova 1939P63
Monilospora (Hacquebard & Barss) Staplin 1960P63
|--*M. moniliformis Hacquebard & Barss 1957 [incl. Densosporites subserratus]P63
|--M. dignata Playford 1963P63
|--M. mutabilis Staplin 1960P63
`--M. triungensis Playford 1963P63
*Reinschospora speciosaP63
Procoronaspora Butterworth & Williams 1958P63
Florinites perlucidus [=Endosporites perlucidus]P63
Paracalathiops [Cycadofilices]P63
`--P. stachei (Stur) Remy 1953P63
Psophosphaera Naumova in Bolkhovitina 1959P63
Aumancisporites Alpern 1958P63
Undulatasporites Leschik 1955P63
Acanthozonotriletes senticosus Ishchenko 1956P63
|--P. glaberP63
|--P. labiatusP63
|--P. parvivermiculatusP63
|--P. pseudobesusP63
`--P. stabilisP63
Phyllothecotriletes rigidusP63
|--V. eximiusP63
`--V. gobbettiiP63
Lophotriletes coniferus Hughes & Playford 1961P63
Apiculatisporis macrurusP16
|--A. mirus Ishchenko 1956P63
`--A. multisetusP63
|--C. clavata (Ishchenko) Hughes & Playford 1961P63
|--C. crassaP63
|--C. harlandiiP63
|--C. labiataP63
|--C. tuberculata (Waltz) Hoffmeister, Staplin & Malloy 1955P63
|--C. usitataP63
`--C. vermiformis Hughes & Playford 1961P63
|--S. clarus Ishchenko 1958P63
|--S. facilisP63
| |--S. f. var. facilisP63
| `--S. f. var. praecrassusP63
|--S. inductus Ishchenko 1956P63
|--S. perforatusP63
|--S. simplexP63
|--S. spetcandusP63
`--S. stenozonalis (Waltz) Ishchenko 1958P63
|--W. albertensis Staplin 1960P63
|--W. lobophora (Waltz) Staplin 1960P63
`--W. sagittataP63
Habrothamnus newelliiT-W89
Uropedium lindeniiT-W89
Caleya minorM89
Vertebraria australisSt89
Aeolanthus repensJ54
Schizogyne sericeaMP05
Dicheiranthus plocamoidesMP05
Coronaria flos-cuculiW17
Henophyton desertiD17
Cutandia memphiticaD17
Haplophyllum tuberculatumD17

Nomen nudum: Chrysodium doriae Squinabol 1889Sq89
Cladomnium crenato-obtusum Dusén 1903D03

Anulatisporites orbiculatus (Waltz) Playford 1963 [=Zonotriletes orbiculatus Waltz in Luber & Waltz 1941, Euryzonotriletes orbiculatus]P63

Colea ramiflora DC. 1845 [=Bignonia ramiflora Decne 1834; incl. B. colei Don 1838, Colea colei Green 1826, C. timorensis ms]S77

Labiadensites fimbriatus (Waltz) Hacquebard & Barss 1957 [=Zonotriletes fimbriatus Waltz in Luber & Waltz 1938, Densosporites fimbriatus (Waltz) Potonié & Kremp 1956, Hymenozonotriletes fimbriatus (Waltz) Ishchenko 1956]P63

Lophozonotriletes rarituberculatus (Luber) Kedo 1957 [=Zonotriletes rarituberculatus Luber in Luber & Waltz 1941; incl. Euryzonotriletes rarituberculatus var. triangulatus Ishchenko 1956, Lophozonotriletes triangulatus Hughes & Playford 1961]P63

*Type species of generic name indicated


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