Embryophyta

Model of the Early Devonian Horneophyton lignieri, by Stephen Caine.


Belongs within: Streptophyta.
Contains: Apo-Tracheophyta, Anthocerotophyta, Marchantiophyta, Bryophyta.

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): Ciliated basal bodies, when present, with distinctive multilayered structure of microtubules and cytoskeletal anchor; open mitosis with phragmoplast at cytokinesis; plasmodesmata and other characteristic cell-cell junctions; diplobiontic life cycle, with vegetative propagation possible in many; alternation of generations, with fertilization of egg by sperm inside protective test; embryology with tissue differentiation coordinated by hormones; differentiated sperm and egg cells, may be on different sexual individuals, on different organs of the same individual, or in the same organ.

Embryophyta [Cormophyta, Embryobiotes, Hepatophyta]
    |--Pan-Tracheophyta [Polysporangiomorpha] CD07
    |    |--Horneophyton lignieri D98, BaCr98
    |    `--Apo-Tracheophyta WM14
    `--+--Anthocerotophyta WM14
       `--+--Marchantiophyta WM14
          `--Bryophyta WM14

Embryophyta incertae sedis:
  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
  Saponaria C55b
    |--S. officinalis C55b
    `--S. vaccaria C55b
  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
  Sabbatia gracilis C55a
  Schultesia C55a
    |--S. brachyptera C55a
    `--S. stenophylla C55b
  Voyria uniflora C55a
  Micranthemum orbiculatum C55a
  Hugueninia tanacetifolia C55a
  Adenostyles leucophylla C55a
  Phyteuma KIW98
    |--P. nigrum KIW98
    |--P. pauciflorum C55a
    `--P. scheuchzeri 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
  Chelidonium majus C55b
  Crambe C55b
    |--C. hispanica C55b
    |--C. maritima C55a
    `--C. orientalis C55b
  Senebiera C55b
    |--S. coronopus C55b
    `--S. pinnatifida C55b
  Cochlearia G60
    |--C. anglica G60
    |--C. danica G60
    |--C. officinalis G60
    `--C. rusticana [incl. C. armoracia] C55b
  Cucubalus bacciferus C55b
  Choerophyllum C55b
    |--C. aromaticum C55b
    `--C. aureum C55b
  Imperatoria ostruthium [=Peucedanum ostruthium] C55b
  Caucalis daucoides 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
  Jussiaea C55b
    |--J. acuminata C55b
    |--J. erecta C55b
    `--J. grandiflora C55b
  Collomia D03
    |--C. gracilis D03
    `--C. grandiflora C55b
  Ervum C55b
    |--E. hirsutum C55b
    `--E. lens C55b
  Helosciadum C55b
    |--H. leptophyllum C55a
    `--H. nodiflorum C55b
  Cnicus benedictus 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
  Leonotis nepetifolia 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
  Paullinia pinnata C55b
  Riedleia concatenata C55b
  Biophytum sensitivum C55b
  Stachytarpha indica C55b
  Centunculus minimus C55b
  Helopus annulatus [=Eriochloa annulata] C55b
  Buchenavia capitata SWK87
  Petitia domingensis SWK87
  Tetrazygia elaeagnoides SWK87
  Exothea paniculata SWK87
  Matayba domingensis SWK87
  Laetia procera SWK87
  Oxandra laurifolia SWK87
  Torrubia fragrans SWK87
  Byrsonia spicata SWK87
  Triphasia trifolia SWK87
  Ventarura lyonii FT05
  Godetia D51
    |--G. amoena D37
    `--G. whitneyi D51
  Penstemon D37
    |--P. azureus D37
    |--P. laetus D37
    `--P. neotericus D37
  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
  Rhoeo discolor D51
  Linanthus parryae D51
  Zauschneria D51
    |--Z. cana D51
    `--Z. septentrionalis D51
  Paederota bonarota PS98
  Physoplexis comosa PS98
  Oglifa gallica BrCa98
  Procopiana insularis S98
  Gymnocarpos decander AGF98
  Anabasis articulata AGF98
  Vaccaria hispanica AGF98
  Dinebra retroflexa GR98
  Guiera senegalensis HK98
  Boscia HK98
    |--B. angustifolia HK98
    `--B. senegalensis HK98
  Microchloa indica HK98
  Serenoa repens CS90
  Plectrachne SM90
    |--P. rigidissima G04
    `--P. schinzii SM90
  Leptosema chambersii SM90
  Trichanthera gigantea DP72
  Blighia unijugata DP72
  Vetiveria elongata WF01
  Tapeinia CL85
    |--T. magellanica D03
    `--T. pumila CL85
  Rhabdotarachnoides Haupt 1957 Du07, H57
    `--*R. simoni Haupt 1957 H57
  Caia BaCr98
  Albertia Schimper in Voltz 1837 FT93
  Swida australis CK05
  Nanophyton erinaceus TLT98
  Chaleuria cirrosa Andrews et al. 1975 G97
  Oocampsa catheta Andrews, Gensel & Kasper 1975 G97
  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
  Streelispora newportensis W97
  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
  Atropis A27
    |--A. novae-zealandiae A27
    `--A. stricta A27
  Narenga porphorycoma BTT02
  Cardiopteridium spitsbergense ZP86
  Pholiurus incurvus BR65
  Koeppenia Schweitzer 2000 S00
    `--*K. eifeliensis Schweitzer 2000 S00
  Pseudosporochnus nodosus S00
  Protolepidodendron scharianum [incl. Leclerqia complexa] S00
  Sartilmania jabachensis S00
  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
  Berula erecta L06
  Balthasaria schliebenii DL94
  Exoecaria agallocha C78
  Gynotroches axillaris C78
  Teysmanniodendron bogoriense C78
  Intsia bijuga C78
  Perezia D03
    |--P. linearis D03
    |--P. magellanica D03
    `--P. pediculariaefolia D03
  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
  Leuceria salina D03
  Nassauvia serpens D03
  Hebenstretia angolensis B06
  Monopsis stellarioides B06
    |--M. s. ssp. stellarioides B06
    `--M. s. ssp. schimperiana 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
  Flourensia cernua MB86
  Symphiocarpos B75
    |--S. occidentalis B75
    `--S. orbiculatus B75
  Drepanostachyum intermedium Da07
  Maotia puya Da07
  Alcimandra cathcartii Da07
  Enhydra fluctuans KLG05
  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
  Aubrieta deltoidea H09
    |--A. d. ssp. deltoidea H09
    `--A. d. ssp. falcata H09
  Hepatica transsilvanica H09
  Calligonum comosum HF03
  Pituranthus triradiatus HF03
  Abronia P81
    |--A. latifolia P81
    |--A. maritima P81
    `--A. umbellata P81
  Dubautia linearis MK08
  Fagaropsis angolensis K09
  Chionodoxa forbesii V09
  Piptochaetium palustre LBM08
  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
  Medeola virginiana A16
  Aloysia wrightii BT72
  Heterotheca subaxillaris BT72
  Hymenothrix wislezeni BT72
  Allionia incarnata BT72
  Hymenopappus lugens BT72
  Oxybaphus comatus BT72
  Antenostoma fasciculatum BT72
  Brickiella californica BT72
  Gaura coccinea BT72
  Cercidium floridum BTA75
  Parmentiera alata BTA75
  Lagascea angustifolia BTA75
  Allenrolfia occidentalis BTA75
  Bouchea prismatica BTA75
  Grindelia BTA75
    |--G. aphanactis BTA75
    `--G. squarosa R35
  Diplopeltis stuartii MLP09
  Carrichtera annua K10
  Hammada L98
    |--H. salicornica L98
    `--H. scoparia L98
  Anvillea garcini L98
  Rhacophyllum coriaceum R87
  Euodia micrococca M87
  Noeggerathiopsis S87
    |--N. media S87
    |--N. prisca S87
    `--N. spathulata S87
  Fortunella japonica B00
  Acradenia euodiiformis B00
  Armeniaca vulgaris TW07
  Peritoma serrulatum R35

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

*Type species of generic name indicated

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