Medullosales

Frond of Neuropteris, copyright Gunnar Ries.

Belongs within: Lignophyta.

The Medullosales are a group of seed ferns known from the late Early Carboniferous to the Permian, generally reconstructed with an unbranching trunk crowned by fronds. At least some genera demonstrate dimorphism in leaf structure, with orbicular 'Cyclopteris' leaves borne on basal parts of the trunk giving way to the pinnate fronds at the crown (Shute & Cleal 2002).

<==Medullosales
    |--Cyclopteris Brongniart 1828 SC02
    |--Odontopteris Brongniart 1828 SC02
    |--Reticulopteris muensteri SC02
    |--Lescuropteris genuina SC02
    |--Blanzyopteris praedentata SC02
    |--Quaestora FC07
    |--Macroneuropteris SC02
    |    |--M. macrophylla SC02
    |    `--M. scheuchzeri SC02
    |--Alethopteris SC02
    |    |--A. decurrens SC02
    |    `--A. lonchitica A38
    |--Laveineopteris Cleal et al. 1990 SC02
    |    |--L. loshii SC02
    |    |--‘Cyclopteris’ orbicularis Brongniart 1831 SC02
    |    |--L. piesbergensis SC02
    |    |--L. rarinervis SC02
    |    `--L. tenuifolia SC02
    `--Neuropteris Brongniart 1828 SC02
         |--N. eriana A38
         |--N. flexuosa R79
         |--N. gigantea ZP86
         |--N. heterophylla (Brongniart) Sternberg 1825 SC02
         |--N. obliqua SC02
         |--N. ovata SC02
         |--N. praedentata H75
         |--N. rarinervis R79
         |--N. selwyni [incl. N. schlehani] A38
         |--N. shaoguangensis ZP86
         `--N. tenuifolia R79

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1–65.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549–552.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2^nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[R79] Ross, C. A. 1979. Carboniferous. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A254–A290. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[SC02] Shute, C. H., & C. J. Cleal. 2002. Ecology and growth habit of Laveineopteris: a gymnosperm from the Late Carboniferous tropical rain forests. Palaeontology 45 (5): 943–972.

[ZP86] Zhang, R., & J. Pojeta Jr. 1986. New bivalves from the Datang Stage, Lower Carboniferous, Guangdong Province, China. Journal of Paleontology 60 (3): 669–679.

Trimerophytina

Life reconstruction of Psilophyton, by Elfriede Abbe.

Belongs within: Tracheophyta.

The Trimerophytina are a group of vascular plants known from the Early (Pragian) to the lowermost Middle Devonian. They bore sporangia in pairs that are commonly preserved twisted around each other (Gerrienne 1997).

See also: Before the word for world was forest.

Characters (from Gerrienne 1997): Branching of axes complex but primarily dichotomous. Sporangia elongate, fusiform; borne terminally in pairs on main or lateral axes, commonly twisted around each other (possibly a standard part of their ontogeny); single longitudinal dehiscence line present, dehiscent sporangia opening into single C-shaped valve.

<==Trimerophytina [Trimerophytophyta, Trimerophytopsida]
    `--Trimerophytaceae G97
         |--Trimerophyton robustius G97
         `--Psilophyton Dawson 1859 G97
              |--P. burnotense G97
              |--P. charientos G97
              |--P. coniculum Trant & Gensel 1985 G97
              |--P. crenulatum G97
              |--P. dapsile Kasper et al. 1974 G97
              |--P. dawsonii Banks et al. 1975 G97
              |--P. forbesii Andrews et al. 1968 G97
              |--P. genseliae Gerrienne 1997 G97
              |--P. krauselii Obrhel 1959 G97
              |--P. microspinosum G97
              |--P. parvulum Gerrienne 1995 G97
              |--P. princeps G97
              `--P. szaferi Zdebska 1986 G97

Trimerophytina incertae sedis:
  Dawsonites G97
    |--D. arcuatus G97
    |--D. magnus G97
    |--D. roskiliensis Chaloner 1972 G97
    `--D. subarcuatus G97
  Hedeia Cookson 1935 G97
    `--H. corymbosa EE86
  Pertica G97
    |--P. dalhousii G97
    |--P. quadrifaria G97
    `--P. varia G97
  Gothanophyton Remy & Hass 1986 G97
  Tursuidea paniculata G97

*Type species of generic name indicated

REFERENCES

[EE86] Edwards, D., & D. S. Edwards. 1986. A reconsideration of the Rhyniophytina Banks. In: Spicer, R. A., & B. A. Thomas (eds)Systematic and Taxonomic Approaches in Palaeobotany.  Systematics Association Special Volume 31: 199–220. Clarendon Press: Oxford.

[G97] Gerrienne, P. 1997. The fossil plants from the Lower Devonian of Marchin (northern margin of Dinant Synclinorium, Belgium): V. Psilophyton genseliae sp. nov., with hypotheses on the origin of Trimerophytina. Review of Palaeobotany and Palynology 98: 303–324.

Dicranophyllales

Reconstruction of Dicranophyllum hallei, from here.

Belongs within: Coniferophyta.

The Dicranophyllales are a group of arborescent plants known from the Carboniferous and Permian, possibly related to modern coniferophytes. They are defined by the having stomata confined to sunken furrows on the abaxial side of the leaf (Rothwell & Mapes 2001).

See also: The Dicranophyllales: an early branch of the conifers?

<==Dicranophyllales
    |--Slivkovia RM01
    |    |--S. lorata RM01
    |    `--S. petschorensis RM01
    |--Entosovia RM01
    |    |--E. kungurica RM01
    |    `--E. rarisulcata RM01
    |--Mostotchkia RM01
    |    |--M. gomankovii RM01
    |    `--M. longifolia RM01
    `--Dicranophyllum RM01
         |--D. effusum RM01
         |--D. gallicum RM01
         |--D. glabrum A38
         |--D. hallei RM01
         |--D. longifolium RM01
         `--D. striatum RM01

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1–65.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637–667.

Miroviaceae

Preserved leaf sections of Mirovia gothanii, from Gomez (2002).

Belongs within: Pinopsida.

The Miroviaceae is an extinct group of conifers known from the mid-Jurassic to the Late Cretaceous, with linear leaves similar in appearance to the modern umbrella pine Sciadopitys verticillata. They differ from modern conifers in having the closely placed stomata restricted to a sunken groove with a protruding margin on the abaxial side of the leaf (Gomez 2002).

<==Miroviaceae
    |--Tritaenia G02
    |--Holkopitys Bose & Manum 1990 G02
    |    `--H. hoegii G02
    |--Oswaldheeria Bose & Manum 1990 G02
    |    |--O. arctica G02
    |    |--O. hallei G02
    |    |--O. macrophylla G02
    |    `--O. scotica G02
    |--Sciadopityoides G02
    |    |--S. crameri G02
    |    |--S. greenboana G02
    |    |--S. ikorfatensis G02
    |    |--S. microphylla G02
    |    |--S. nathorstii G02
    |    |--S. ukrainensis G02
    |    |--S. uralensis G02
    |    `--S. variabilis G02
    `--Mirovia G02
         |--*M. szaferi Reymanówna 1985 G02
         |--M. capbohemanensis G02
         |--M. florinii G02
         |--M. gothanii Gomez 2002 G02
         |--M. groenlandica G02
         |--M. ineffecta G02
         |--M. lagerheimii G02
         |--M. persulcata G02
         `--M. sibirica G02

*Type species of generic name indicated

REFERENCES

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761–773.

Cheirolepidiaceae

Frenelopsis hoheneggeri, from the Peabody Museum of Natural History.

Belongs within: Pinopsida.

The Cheirolepidiaceae are a group of fossil conifers known from the Mesozoic. They are characterised by a distinct pollen type (assigned to the form genus Classopollis) with a complex, tectate-columellate wall structure (Axsmith et al. 2004). Shoots were often opposite-decussate or spiral, and leaves (which may be dimorphic) were typically reduced or scale-like, and imbricate (Pole 2000).

<==Cheirolepidiaceae [Frenelopsidaceae]
    |--Alvinia bohemica AKW04
    |--Tomaxellia biforme P00
    |--Otwayia Pole 2000 P00
    |    |--*O. tetragona (Cantrill & Douglas) Pole 2000 [=Geinitzea tetragona Cantrill & Douglas 1988] P00
    |    `--O. cudgeloides Pole 2000 P00
    |--Classostrobus Alvin, Spicer & Watson 1978 AKW04 [incl. Pseudofrenelopsis AKW04, Suturovagina GM-C02]
    |    |--*C. comptonensis Alvin, Spicer & Watson 1994 AKW04 (see below for synonymy)
    |    |--C. arkansensis Axsmith, Krings & Waselkov 2004 AKW04
    |    |--C. cathayanus [incl. Pseudofrenelopsis intermedia] AKW04
    |    |--C. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
    |    `--C. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
    |--Classopollis Pflug 1953 AKW04
    |    |--*C. classoides Pflug 1953 GM-C02
    |    |--C. brasiliensis Herngreen 1975 I02
    |    |--C. mirabilis Reyre 1970 GM-C02
    |    |--C. noeli Reyre 1970 GM-C02
    |    `--C. obidosensis Groot & Groot 1962 GM-C02
    |--Cupressinocladus Seward 1919 AKW04 [incl. Watsoniocladus GM-C02]
    |    |--C. asferensis Barale & Ouaja 2002 BO02
    |    |--C. itieri (Saporta) Barale 1981 BO02
    |    |--C. leptocladoides (Berry) Pons 1990 BO02
    |    |--C. micromerum (Heer) Pais 1974 BO02
    |    |--C. pompeckji (Salfeld) Pons 1990 BO02
    |    |--C. pseudoexpansum Barnard & Miller 1976 BO02
    |    `--C. soltanensis Barale, Ouaja & Philippe 2000 B)2
    `--Frenelopsis Schenck 1869 AKW04, GM-C02
         |--*F. hoheneggeri (Ettingshausen) Schenck 1869 [=Thuites hoheneggeri Ettinghausen 1852] GM-C02
         |--F. alata Feistmantel 1881 AKW04
         |--F. choshiensis GM-C02
         |--F. elegans GM-C02
         |--F. harrisii Doludenko & Reymanówna 1978 GT02
         |--F. kaneviensis Barale & Doludenko 1985 GM-C02
         |--F. occidentalis GM-C02
         |--F. oligostomata AKW04 [incl. Classoidites glandis GM-C02]
         |--F. pombetsuensis Saiki 1997 GM-C02
         |--F. ramosissima GM-C02
         |--F. rubiesensis Barale 1973 GM-C02
         |--F. sifloana Watson 1983 GM-C02
         |--F. teixeirae GM-C02
         |--F. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
         |--F. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
         `--F. veneta Gomez, Thévenard et al. 2002 GT02

*Classostrobus comptonensis Alvin, Spicer & Watson 1994 AKW04 [incl. Classopollis martinottii GM-C02, Pseudofrenelopsis parceramosa (Fontaine) Watson 1977 AKW04]

*Type species of generic name indicated

REFERENCES

[AKW04] Axsmith, B. J., M. Krings & K. Waselkov. 2004. Conifer pollen cones from the Cretaceous of Arkansas: implications for diversity and reproduction in the Cheirolepidiaceae. Journal of Paleontology 78: 402–409.

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

[GM-C02] Gomez, B., C. Martín-Closas, G. Barale, N. Solé de Porta, F. Thévenard & G. Guignard. 2002. Frenelopsis (Coniferales: Cheirolepidiaceae) and related male organ genera from the Lower Cretaceous of Spain. Palaeontology 45 (5): 997–1036.

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775–788.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

Lignophyta

Reconstruction of Archaeopteris, from here.

Belongs within: Tracheophyta.
Contains: Medullosales, Spermatophyta.

The Lignophyta comprises the clade of wood-producing plants, defined by Cantino et al. (2007) as the most inclusive clade characterised by a bifascial vascular cambium (a meristematic layer producing secondary xylem on the inner face and secondary phloem on the outer face) homologous with that of Pinus sylvestris. Synapomorphies of lignophytes also include a periderm-producing cork cambium, and cortical fibre strands (lost in most derived subgroups; Cantino et al. 2007).

<==Lignophyta (see below for synonymy)
    |--Aneurophytales CD07
    |--Tetraxylopteris FC07
    `--+--+--Cecropteris FC07
       |  `--Archaeopteris Dawson 1871 FC07, C81 [Archaeopteridales]
       |       |--*A. hibernica C81, CWC87 [=Cyclopteris hibernicus Forbes 1854 C81]
       |       |--A. changyangensis CWC87
       |       |--A. fimbricata C81
       |       |--A. fissilis CWC87
       |       |--A. gothani [=Adiantites gothani] C81
       |       |--A. halliana CWC87
       |       |--A. howitti F71
       |       |--A. hubeiensis CWC87
       |       |--A. jacksoni CWC87
       |       |--A. latifolia C81
       |       |--A. macilenta CWC87
       |       |--A. mapaensis [=Cardiopteridium mapaense] C81
       |       |--A. obtusa CWC87
       |       |--A. roemeriana (Goeppert) Lesquereux 1880 (see below for synonymy) CWC87
       |       |--A. rogersi CWC87
       |       |--A. sphenophyllifolia CWC87
       |       `--A. wilkinsoni Feistmantel 1878-1879 F71
       `--+--Elkinsia polymorpha FC07, MM12
          `--+--Laceya FC07
             `--+--+--Bilignea FC07
                |  `--Lyrasperma FC07
                `--Apo-Spermatophyta [Gymnospermae, Spermatophytata] CD07
                     |  i. s.: Dicksonites pluckeneti T30
                     |         Rienitsia variabilis M01
                     |--Heterangium FC07
                     |--+--Medullosales FC07
                     |  `--+--Callistophyton [Callistophytaceae] FC07
                     |     |    `--C. proxyloides RMH-C05
                     |     `--Spermatophyta FC07
                     `--Lyginopteridaceae D98
                          |--Lyginopteris FC07
                          |--Pecopteris VA01
                          |    |--P. obscura Dun 1897 F71
                          |    `--P. plumosa A38
                          |--Rhacopteris VA01
                          |    |--R. fertilis VA01
                          |    |--R. intermedia F71
                          |    |--R. lindsaeformis VA01
                          |    |--R. paniculifera VA01
                          |    `--R. septentrionalis Feistmantel 1878-1879 F71
                          `--Sphenopteris VA01
                               |--S. bifida VA01
                               |--S. eocenica S89
                               |--S. grandis Dun 1898 F71
                               |--S. marginata [incl. S. rotundifolia] A38
                               |--S. obtusiloba CWC87
                               `--S. valida [incl. S. artemisiae-folioides] A38

Lignophyta incertae sedis:
  Botrychiopsis Z02
  Genselia uberis VA01
  Triphyllopteris [Calamopityaceae] VA01
  Alisporites YB02
    |--A. bilateralis Rouse 1959 YB02
    `--A. similis (Balme) Dettmann 1963 YB02
  Ephedripites YB02
    |--E. eocaenicus (Selling) Yi & Batten 2002 [=Schizaea eocaenica Selling 1944] YB02
    |--E. huanghuaensis Ke & Shi 1978 YB02
    |--E. irregularis Herngreen 1973 I02
    |--E. jansonii (Pocock) Muller 1968 YB02
    |--E. multicostatus Brenner 1963 YB02
    |--E. multipartitus (Chlonova) Gao & Zhao 1976 YB02
    |--E. ovalis Muller 1968 I02
    `--E. praeclarus (Chlonova) Yi & Batten 2002 [=Schizaea praeclara Chlonova 1961] YB02
  Inaperturopollenites YB02
    |--I. dubius (Potonié & Venitz) Thomson & Pflug 1953 YB02
    |--I. hiatus (Potonié) Thomson & Pflug 1953 YB02
    `--I. laevigatus Takahashi 1957 YB02
  Cedripites parvus Norton in Norton & Hall 1969 YB02
  Phyllocladidites YB02
    |--P. inchoatus (Pierce) Norris 1967 YB02
    `--P. mawsonii Cookson 1947 RC02
  Pinuspollenites labdacus YB02
    |--P. l. f. labdacus YB02
    `--P. l. f. maximus Potonié 1958 YB02
  Podocarpidites YB02
    |--P. minutus Wang 1975 YB02
    `--P. nageiaformis (Zaklinskaya) Takahashi 1964 YB02
  Rugubivesiculites YB02
    |--R. convolutus Pierce 1961 YB02
    |--R. reductus Pierce 1961 YB02
    `--P. rugosus Pierce 1961 YB02
  Protopityales CD07
  Crossia SS04

Archaeopteris roemeriana (Goeppert) Lesquereux 1880 [=Cyclopteris roemeriana Goeppert 1859; incl. A. guangdongensis Feng 1977, A. mutatoformis Li 1984, A. tonglingiana Li 1984, A. zhongmingiana Li 1984] CWC87

Lignophyta [Cycadophytanae, Lignophytia, Pan-Spermatophyta, Progymnospermida, Progymnospermophyta, Progymnospermopsida, Radiatopses]

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1-65.

[C81] Cai C.-Y. 1981. On the occurrence of Archaeopteris in China. Acta Palaeontologica Sinica 20 (1): 75-80.

[CWC87] Cai C.-Y., Wen Y.-G. & Chen P.-Q. 1987. Archaeopteris florula from Upper Devonian of Xinhui County, central Guangdong and its stratigraphical significance. Acta Palaeontologica Sinica 26 (1): 55-64.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549-552.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775-788.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793-7798.

[M01] McLoughlin, S. 2001. The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany 49: 271-300.

[RC02] Riding, J. B., & J. A. Crame. 2002. Aptian to Coniacian (Early–Late Cretaceous) palynostratigraphy of the Gustav Group, James Ross Basin, Antarctica. Cretaceous Research 23: 739-760.

[RMH-C05] Rothwell, G. W., G. Mapes & G. R. Hernandez-Castillo. 2005. Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54 (3): 733-750.

[SS04] Schneider, H., E. Schuettpelz, K. M. Pryer, R. Cranfill, S. Magallón & R. Lupia. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553-557.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 73-76.

[T30] Thomas, H. H. 1930. Further observations on the cuticle structure of Mesozoic cycadean fronds. Proceedings of the Linnean Society of London 141: 5-7.

[VA01] Vega, J. C., & S. Archangelsky. 2001. Austrocalyxaceae, a new pteridosperm family from Gondwana. Palaeontographica Abt. B 257: 1-16.

[YB02] Yi, S., & D. J. Batten. 2002. Palynology of Upper Cretaceous (uppermost Campanian-Maastrichtian) deposits in the South Yellow Sea Basin, offshore Korea. Cretaceous Research 23: 687-706.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

Tracheophyta

Cooksonia pertonii, from here.

Belongs within: Apo-Tracheophyta.
Contains: Trimerophytina, Lignophyta, Monilophyta, Lycopodiopsida.

Tracheophyta was defined by Cantino et al. (2007) as the crown-group of vascular plants, i.e. the largest crown clade including Zea mays but not Phaeoceros laevis, Marchantia polymorpha or Polytrichum commune, characterised by the walls of the water-conducting cells having a thick, lignified, decay-resistent layer. Modern tracheophytes are divided between the Lycopodiopsida (clubmosses and allies) and Euphyllophyta (ferns and seed plants). Living members of the latter clade are characterised by the presence of broad true leaves (megaphylls or euphylls), but fossil evidence suggests that the leaves of ferns and seed plants may have evolved independently from each other (Cantino et al. 2007).

Tracheophyta [Cormatae, Pteridophytina]
    |--Pan-Euphyllophyta [Euphyllophytina] CD07
    |    |--Trimerophytina CD07
    |    `--Euphyllophyta CD07
    |         |--Lignophyta CD07
    |         `--Pan-Monilophyta [Pteridophytanae] CD07
    |              |--Cladoxylopsida BC98
    |              |    |--Hyeniales D98
    |              |    `--Cladoxylon [Cladoxylales] D98
    |              |         `--C. dawsonii D98
    |              `--+--Monilophyta CD07
    |                 |--Rhacophyton D98
    |                 `--Zygopteris [Zygopteridales] D98
    `--Pan-Lycopodiophyta [Lycopodiobiotina] CD07
         |--Cooksonia Lang 1937 D98, EE86
         |    |--C. bohemica EE86
         |    |--C. caledonica EE86
         |    |--C. crassiparietilis EE86
         |    |--C. hemisphaerica EE86
         |    `--C. pertonii BC98
         `--+--Renalia D98
            |    |--R. grabertii EE86
            |    |--R. hueberi EE86
            |    `--R. major EE86
            `--Lycophytina BC98
                 |--Lycopodiopsida CD07
                 `--Zosterophyllopsida BC98
                      |--Gosslingia breconensis BC98
                      |--Zosterophyllum rhenanum D98, S00
                      `--Sawdonia D98
                           |--S. ornata S00
                           `--S. spinosissima S00

Tracheophyta incertae sedis:
  Carpolithes CWC87
    |--C. amaranthaceus Ettingshausen 1886 F71
    |--C. heerii JD05
    `--C. wetherellioides Ettingshausen 1886 F71
  Xylia PA-W02
    |--X. dolabriformis PA-W02
    `--X. xylocarpa RH91
  Paripteris pseudogigantea SC02
  Austrocalyxaceae VA01
    |--Rinconadia archangelskyi VA01
    |--Jejenia alata VA01
    |--Austrocalyx Vega & Archangelsky 1996 VA01
    |    `--A. jejenensis VA01
    `--Polycalyx Vega & Archangelsky 2001 VA01
         |--*P. laterale Vega & Archangelsky 2001 VA01
         `--P. tetramera Vega & Archangelsky 2001 VA01
  Nothorhacopteris VA01
  Bergiopteris VA01
  Hydrasperma tenuis VA01
  Diplopteridium VA01
    |--D. holdenii VA01
    `--D. teilianum VA01
  Pullaritheca longii [=Hydrasperma longii] VA01
  Moresnetia VA01
  Buteoxylonaceae VA01
  Stauropteridales BC98
  Yunia dichotoma Hao & Beck 1991 G97
  Lycopodiella serpentina P05
  Cicatricosisporites I02
    |--C. annulatus VB02
    |--C. augustus VB02
    |--C. australiensis (Cookson) Potonié 1956 I02
    |--C. hughesii VB02
    |--C. ludbrookiae VB02
    |--C. orbiculatus Singh 1964 I02
    |--C. purbeckensis VB02
    `--C. recticicatricosus Döring 1965 I02
  Deltoidospora I02
    |--D. hallii Miner 1935 I02
    `--D. psilostoma Rouse 1959 YB02
  Gleicheniidites senonicus Ross 1949 I02
  Matonisporites cooksoniae I02, VB02
  Cibotiumspora jurienensis (Balme) Filatoff 1975 I02
  Cyathidites I02
    |--C. australis Couper 1953 YB02
    `--C. minor Couper 1953 YB02
  Todisporites I02
    |--T. major Couper 1958 YB02
    `--T. minor Couper 1958 YB02
  Trilobosporites laevigatus El Beialy 1994 I02
  Camelina sativa BO08
  Aneimites F71
    |--A. austrina Etheridge 1888 F71
    `--A. ovata [incl. Rhacopteris inaequilatera] F71
  Apocynocarpum sulcatum Ettingshausen 1886 F71
  Artocarpidium F71
    |--A. gregoi Ettingshausen 1886 F71
    `--A. stuarti Ettingshausen 1883 F71
  Bairera multifida F71
  Bambusites arthrostylinus Ettingshausen 1886 F71
  Blechnoxylon talbragarense Etheridge 1899 F71
  Walkomiella australis [=Brachyphyllum australe Feistmantel 1878-1879] F71
  Callistemophyllum F71
    |--C. hackii Ettingshausen 1886 F71
    `--C. obliquum Ettingshausen 1886 F71
  Cladophlebis F71
    |--C. australis F71
    `--C. roylei F71
         |--C. r. var. roylei F71
         `--C. r. var. murtoni Dun 1909 F71
  Cupanites selwyni Ettingshausen 1886 F71
  Cycadopteris scolopendrina Ratte 1887 F71
  Dalbergiophyllum affine Ettingshausen 1886 F71
  Diemenia perseaefolia Ettingshausen 1886 F71
  Dolichites coriaceus Ettingshausen 1886 F71
  Ficonium solandri Ettingshausen 1883 F71
  Gangamopteris walkomii Rigby 1967 F71
  Ginkgocladus australiensis Ettingshausen 1886 F71
  Heterocladiscos thujoides Ettingshausen 1883 F71
  Salisburia palmata [=Jeanpaulia palmata Ratte 1886] F71
  Lepidopteris madagascariensis F71
  Macrotaeniopteris wianamattae Feistmantel 1878-1879 F71
  Molongia brownii Ettingshausen 1883 F71
  Maitaia podocarpoides F71
  Malpighiastrum babbagei Ettingshausen 1886 F71
  Microphyllopteris pectinata F71
  Myrtonium F71
    |--M. lanceolatum Ettingshausen 1886 F71
    `--M. obtusifolium Ettingshausen 1886 F71
  Oleandridium lenticuliforme Etheridge 1894 F71
  Ovopteris alata F71
  Palaeocladus cuneiformis Ettingshausen 1886 F71
  Pitys sussmilchi F71
  Pseudopinus wilkinsoni Ettingshausen 1886 F71
  Rhopala sapindifolia Ettingshausen 1886 F71
  Rissikia F71
    |--R. apiculata Townrow 1967 F71
    `--R. media F71
  Stenopteris rigida Dun 1909 F71
  Thinnfeldia F71
    |--T. feistmanteli F71
    |--T. lancifolia F71
    `--T. narrabeenensis Dun in Walkom 1925 F71
  Dicroidium odontopteroides [=Thinnfeldia odontopteroides] F71
  Trachyphyllum myosotinum Ettingshausen 1886 F71
  Leptophloeum F71
    |--L. australe [=Lepidodendron australe] F71
    `--L. rhombicum ZA02
  Annularia F71
    |--A. australis Feistmantel 1878-1879 F71
    |--A. ivini Walkom 1941 F71
    |--A. latifolia A38
    |--A. sphenophylloides A38
    `--A. stellata A38
  Dryophyllum F71
    |--D. curticellense JD05
    |--D. howitti Ettingshausen 1886 F71
    `--D. moselense JD05
  Cyclostigma F71
    |--C. australe Feistmantel 1878-1879 F71
    `--C. kiltorkense CWC87

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1-65.

[BC98] Bateman, R. M., P. R. Crane, W. A. DiMichele, P. R. Kenrick, N. P. Rowe, T. Speck & W. E. Stein. 1998. Early evolution of land plants: phylogeny, physiology, and ecology of the primary terrestrial radiation. Annual Review of Ecology and Systematics 29: 263-292.

[BO08] Bönsel, D., I. Ottich, A. Malten & G. Zizka. 2008. An updated list of the vascular plants of Frankfurt am Main (Pteridophyta & Spermatophyta). Senckenbergiana Biologica 88 (1): 111-121.

[CWC87] Cai C.-Y., Wen Y.-G. & Chen P.-Q. 1987. Archaeopteris florula from Upper Devonian of Xinhui County, central Guangdong and its stratigraphical significance. Acta Palaeontologica Sinica 26 (1): 55-64.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[EE86] Edwards, D., & D. S. Edwards. 1986. A reconsideration of the Rhyniophytina Banks. In Systematic and Taxonomic Approaches in Palaeobotany (R. A. Spicer & B. A. Thomas, eds) Systematics Association Special Volume 31: 199-220. Clarendon Press: Oxford.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[G97] Gerrienne, P. 1997. The fossil plants from the Lower Devonian of Marchin (northern margin of Dinant Synclinorium, Belgium): V. Psilophyton genseliae sp. nov., with hypotheses on the origin of Trimerophytina. Review of Palaeobotany and Palynology 98: 303-324.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775-788.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185-273.

[PA-W02] Pang, K.-L., M. A. Abdel-Wahab, S. Sivichai, H. M. El-Sharouney & E. B. G. Jones. 2002. Jahnulales (Dothideomycetes, Ascomycota): A new order of lignicolous freshwater ascomycetes. Mycological Research 106 (9): 1031-1042.

[P05] Pinder, A. M. 2005. A review of biodiversity in wetlands with organic sediments on the Swan Coastal Plain, Western Australia, with an emphasis on aquatic invertebrates. Journal of the Royal Society of Western Australia 88 (3): 129-132.

[RH91] Ramani, N., & M. A. Haq. 1991. Potential of Meristacarus degradatus and Xylobates rhomboides (Acari: Oribatei) in the degradation of higher plant materials. In Dusbábek, F. & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 411-415. SPB Academic Publishing: The Hague.

[S00] Schweitzer, H. J. 2000. Neue Pflanzenfunde im Unterdevon der Eifel (Deutschland). Senckenbergiana Lethaea 80 (2): 371-395.

[SC02] Shute, C. H., & C. J. Cleal. 2002. Ecology and growth habit of Laveineopteris: A gymnosperm from the Late Carboniferous tropical rain forests. Palaeontology 45 (5): 943-972.

[VA01] Vega, J. C., & S. Archangelsky. 2001. Austrocalyxaceae, a new pteridosperm family from Gondwana. Palaeontographica Abt. B 257: 1-16.

[VB02] Vijaya & T. K. Bhattacharji. 2002. An Early Cretaceous age for the Rajmahal traps, Panagarh area, West Bengal: Palynological evidence. Cretaceous Research 23: 789-805.

[YB02] Yi, S., & D. J. Batten. 2002. Palynology of Upper Cretaceous (uppermost Campanian-Maastrichtian) deposits in the South Yellow Sea Basin, offshore Korea. Cretaceous Research 23: 687-706.

[ZA02] Zhu, M., P. E. Ahlberg, W. Zhao & L. Jia. 2002. First Devonian tetrapod from Asia. Nature 420: 760-761.

Coniferophyta

Maidenhair tree Ginkgo biloba, copyright Liné1.

Belongs within: Spermatophyta.
Contains: Dicranophyllales, Pinopsida.

The Coniferophyta has been recognised as a clade of gymnosperms including conifers, the ginkgo Ginkgo biloba, and the Palaeozoic Cordaitales. Members of this clade have simple, dichotomously veined leaves, simple strobili in the axils of the leaves, and platyspermic seeds (Doyle 1998).

<==Coniferophyta
    |  i. s.: Carpentieria frondosa RM01
    |         Dichophyllum RM01
    |         Ginkgophyllum RM01
    |         Polyspermophyllum sergii RM01
    |         Sphenobaiera RM01
    |         Vojnovskyales RM01
    |           |--Sergeia neuburgii RM01
    |           `--Vojnovskya [Vojnovskyaceae] RM01
    |                `--V. paradoxa RM01
    |--Ginkgoopsida [Ginkgophyta] RY03
    |    |  i. s.: Ginkgoites RY03
    |    |         Phoenicopsis RY03
    |    |         Trichopitys heteromorpha RM01
    |    |         Karkenia D98
    |    |         Baiera D98
    |    `--Ginkgoales FC07
    |         |--Nehvizdya penalveri G02
    |         |--Ginkgo [Ginconinae, Ginkgoideae] CD07
    |         |    |--G. baiera GT12
    |         |    |--G. biloba Linnaeus 1771 CD07
    |         |    |--G. huttoni PL12
    |         |    `--G. liaoningensis PL12
    |         |--Eretmophyllum PL12
    |         |--Pseudotorellia PL12
    |         `--Nehvizdyella bipartita PL12
    `--Pan-Coniferae CD07
         |--+--Dicranophyllales RMH-C05
         |  `--Pinopsida FC07
         `--Cordaitales [Cordaitanthales] FC07
              |  i. s.: Cordaites F71
              |           |--C. australis F71
              |           |--C. borassifolius [=Flabellaria borassifolia] F04
              |           `--C. robbii [incl. C. borassifolius] A38
              |         Cordaitanthus devonicus A38
              |--Shanxioxylon FC07
              `--+--Mesoxylon priapii FC07, RMH-C05
                 `--Cordaixylon FC07
                      |--C. dumusum RMH-C05
                      `--C. iowensis MM12

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1-65.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549-552.

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

[GT12] Giribet, G., A. L. Tourinho, C. Shih & D. Ren. 2012. An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Organisms, Diversity & Evolution 12: 51-66.

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761-773.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793-7798.

[PL12] Peñalver, E., C. C. Labandeira, E. Barrón, X. Delclòs, P. Nel, A. Nel, P. Tafforeau & C. Soriano. 2012. Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the USA 109 (22): 8623-8628.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1-11.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637-667.

[RMH-C05] Rothwell, G. W., G. Mapes & G. R. Hernandez-Castillo. 2005. Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54 (3): 733-750.

Pteridaceae

Mangrove fern Acrostichum aureum, photographed by Andrés Hernández.

Belongs within: Pteridales.
Contains: Vittariaceae, Adiantum, Pteris.

The Pteridaceae is a cosmopolitan family of ferns with abaxial sori lacking indusia or protected by a reflexed or revolute leaf margin (M. D. Windham). The molecular phylogenetic analysis of ferns by Schneider et al. (2004) suggested that the Pteridaceae may be paraphyletic with regard to the Vittariaceae. The Pteridaceae also includes the aquatic genus Ceratopteris, as well as the semi-aquatic Acrostichum. Cheilanthes, the lip ferns, is a cosmopolitan genus of ferns found growing on rocks in warm, dry regions.

Characters (from M. D. Windham): Plants perennial (occasionally annual), on rock or terrestrial, of small (rarely large) stature . Stems compact to creeping, branched or unbranched, dictyostelic, bearing hairs and/or scales. Leaves monomorphic to dimorphic, circinate or noncircinate in bud . Petioles usually with persistent scales proximally, lacking spines; vascular bundles 1-several, roundish or crescent-shaped in cross section. Blades 1-6-pinnate, without laminar buds . Indument on petioles, rachises, costae, and blades, rarely absent or commonly of hairs, glands, and/or scales, occasionally of white or yellow farina. Veins pinnate or parallel in ultimate segments of blades, simple or forked, free or infrequently anastomosing in complex patterns. Sori borne abaxially on veins, often confluent with age and forming a continuous submarginal band, or sporangia densely covering abaxial surface (acrostichoid); receptacle not or only slightly elevated. Indusia (when present) formed by reflexed, recurved, or revolute leaf margin (false indusium). Sporangia stalk of 2--3 rows of cells; annulus vertical, interrupted by stalk; spores 64 or 32 (rarely 16) per sporangium. Spores all 1 kind, brown, black, or gray (rarely yellow), globose to globose-tetrahedral or trigonal, occasionally with prominent equatorial ridge, trilete, or trigonal, variously ornamented (usually cristate or rugose). Gametophytes green, aboveground, obcordate to reniform, sometimes asymmetric, usually glabrous (glandular-farinose in Notholaena); archegonia and antheridia borne on abaxial surface, antheridia 3-celled.

Pteridaceae [Adiantaceae]
    |--+--Bommeria SS04
    |  |    |--B. ehrenbergiana SS04
    |  |    `--B. pedata L54
    |  `--+--Vittariaceae SS04
    |     `--+--Adiantum SS04
    |        `--Hewardia regia SS04
    `--+--Pteris SS04
       `--+--Acrostichum SS04
          |    |--A. aureum SS04
          |    |--A. danaeifolium SS04
          |    `--A. speciosum WF01
          `--+--Magnastriatites SS04
             `--Ceratopteris SS04
                  |--C. richardii SL05
                  `--C. thalictroides (Linnaeus) Brongniart 1822 (see below for synonymy) I88

Pteridaceae incertae sedis:
  Argyrochosma YS03
  Anopteris hexagonia J87
  Gymnopteris Bernhardi 1799 YS03
    `--G. vestita (Bedd.) Underwood 1902 (see below for synonymy) I88
  Notholaena Brown 1810 [=Chrysochosma (Kümm.) Pichi Sermolli 1989] YS03
    |--*N. trichomanoides (Linnaeus) Brown 1810 [=Pteris trichomanoides] YS03
    `--N. parryi GCR98
  Paragymnopteris Shing 1993 YS03
    `--*P. marantae (Linnaeus) Shing 1993 (see below for synonymy) YS03
  Pellaea Link 1841 A61
    |--P. atropurpurea SL05
    |--P. falcata (Br.) Fée 1850-1852 (see below for synonymy) A61
    `--P. rotundifolia (Forster) Hook. 1858 (see below for synonymy) A61
  Cheilanthes Swartz 1806 A61 [incl. Aleuritopteris I88]
    |--C. albomarginata Clarke 1880 [=C. farinosa var. albomarginata (Clarke) Bedd. 1892] I88
    |--C. anceps Blanford 1886 I88
    |--C. angustifolia L54
    |--C. aurea L54
    |--C. austrotenuifolia G04
    |--C. brownii LK14
    |--C. caudata LK14
    |--C. chrysophylla Hooker 1852 [=C. farinosa var. chrysophylla (Hooker) Clarke 1880] I88
    |--C. contigua LK14
    |--C. dalhousiae Hooker 1852 (see below for synonymy) I88
    |--C. deltoidea KLN06
    |--C. distans (Brown) Mett. 1859 [=Notholaena distans Brown 1810] A61
    |--C. duthiei Baker 1891 I88
    |--C. farinosa (Forskål) Kaulf. 1824 (see below for synonymy) I88
    |--C. fragillima LK14
    |--C. fragrans D30
    |--C. kaulfussii L54
    |--C. lanosa SL05
    |--C. lasiophylla G04
    |--C. notholaenoides J87
    |--C. nudiuscula LK14
    |--C. odora C74
    |--C. persica GCR98
    |--C. pteridioides PT98
    |--C. pumilio LK14 [=Notholaena pumilio Brown 1810 YS03]
    |--C. pyramidalis L54
    |--C. rufa Don 1825 I88
    |--C. setigera Blume 1828 A61 (see below for synonymy)
    |--C. sieberi Kunze in Lehm. 1846 [incl. C. erecta Colenso 1896] A61
    |--C. subvillosa Hooker 1852 [=Aleuritopteris subvillosa (Hooker) Ching 1941] I88
    |--C. tamburii (Hooker) Moore 1861 [=Pellaea tamburii Hooker 1858] I88
    |--C. tenuifolia (Burmann) Swartz 1806 I88 (see below for synonymy)
    `--C. vellea PT98 [=Notholaena vellea Brown 1810 YS03]
  Neurocallis SS04
  Paesia St. Hilaire 1833 A61
    `--P. scaberula (Rich.) Kuhn 1882 (see below for synonymy) A61
  Paraceterach muelleri LK14
  Platyzoma microphyllum LK14

Nomina nuda: Pellaea saportana Squinabol 1889 S89

Ceratopteris thalictroides (Linnaeus) Brongniart 1822 [=Acrostichum thalictroides Linnaeus 1753; incl. A. siliquosum Linnaeus 1753, Ceratopteris siliquasa (Linnaeus) Copel. 1935] I88

Cheilanthes dalhousiae Hooker 1852 [=C. farinosa var. dalhousiae (Hooker) Clarke 1880, Leptolepidium dalhousiae] I88

Cheilanthes farinosa (Forskål) Kaulf. 1824 [=Pteris farinosa Forskål 1775; incl. C. dealbata Don 1825 non Pursh 1814, C. grisea Blanford 1886, Aleuritopteris grisea (Blanford) Panigr. 1961] I88

Cheilanthes setigera Blume 1828 A61 [incl. Dryopteris setigera A61, Thelypteris setigera (Blume) Ching 1936 C49]

Cheilanthes tenuifolia (Burmann) Swartz 1806 I88 [=Trichomanes tenuifolium Burmann 1768 I88; incl. C. kirkii Armstrong 1881 (n. n.) A61, C. venosa Colenso 1893 A61]

Gymnopteris vestita (Bedd.) Underwood 1902 [=Syngramma vestita Bedd. 1883, Grammitis vestita Wallich ex Bedd. 1883 (n. n.), Gymnogramma vestita] I88

Paesia scaberula (Rich.) Kuhn 1882 [=Pteris scaberula Rich. 1832, Allosurus scaberulus Presl 1836; incl. Pt. microphylla Cunn. 1837] A61

*Paragymnopteris marantae (Linnaeus) Shing 1993 [=Acrostichum marantae Linnaeus 1753, Notholaena marantae (Linnaeus) Brown 1810, Paraceterach marantae (Linnaeus) Tryon 1986] YS03

Pellaea rotundifolia (Forster) Hook. 1858 [=Pteris rotundifolia Forster 1786, Allosurus rotundifolius Kunze 1856, Platyloma rotundifolia Smith 1841] A61

Pteris falcata (Br.) Fée 1850-1852 [=Pteris falcata Br. 1810, Platyloma falcata Smith 1841; incl. Pt. seticaulis Hook. 1840] A61

*Type species of generic name indicated

REFERENCES

[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens). Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1–299, 224 pls.

[C49] Crookes, M. W. 1949. A revised and annotated list of New Zealand Filicinae. Transactions and Proceedings of the Royal Society of New Zealand 77 (2): 209–225.

[D30] Druce, G. C. 1930. Account of a botanical tour in Cyprus. Proceedings of the Linnean Society of London 141: 50–52.

[G04] Gibson, N. 2004. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35–47.

[GCR98] Gratani, L., M. F. Crescente & G. Rossi. 1998. Photosynthetic performance and water use efficiency of the fern Cheilanthes persica. Photosynthetica 35: 507–516.

[I88] Iwatsuki, K. 1988. An enumeration of the pteridophytes of Nepal. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 231–339.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

[KLN06] Klopper, R. R., P. Lemmer & J. Nel. 2006. Pteridophyta: Pteridaceae. Cheilanthes deltoidea, a new locality in Gauteng, South Africa. Bothalia 36 (2): 173–174.

[L54] Lötschert, W. 1954. Beitrag zur Pteridophyten-Flora von Mittel-Amerika. Senckenbergiana Biologica 35 (1–2): 109–119.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[SS04] Schneider, H., E. Schuettpelz, K. M. Pryer, R. Cranfill, S. Magallón & R. Lupia. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553–557.

[SL05] Small, R. L., E. B. Lickey, J. Shaw & W. D. Hauk. 2005. Amplification of noncoding chloroplast DNA for phylogenetic studies in lycophytes and monilophytes with a comparative example of relative phylogenetic utility from Ophioglossaceae. Molecular Phylogenetics and Evolution 36: 509–522.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 73–76.

[WF01] Woinarski, J. C. Z., A. Fisher, K. Brennan, I. Morris & R. Chatto. 2001. Patterns of bird species richness and composition on islands off Arnhem Land, Northern Territory, Australia. Austral Ecology 26: 1–13.

[YS03] Yatskievych, G., & A. R. Smith. 2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52: 331–336.

Cupressaceae

Coastal redwoods Sequoia sempervirens, copyright Acroterion.

Belongs within: Cupressophyta.
Contains: Taxodioideae, Callitroideae, Cupressoideae.

The conifer family Cupressaceae includes cypresses (Cupressus), junipers (Juniperus) and related taxa. The family has previously been divided between the Cupressaceae sensu stricto (with leaves opposite or whorled, comprising Callitroideae and Cupressoideae) and Taxodiaceae (with leaves mostly alternate), but the latter is now recognised as paraphyletic to the former and most recent authors combine the two. Many species are significant timber trees. The giant redwood Sequoiadendron giganteum is the world's largest (though not tallest) tree.

Members of the Cupressaceae include the east Asian genera Cunninghamia and Taiwania, both of which bear seed cones with thin, leathery scales; Cunninghamia has linear, tapering leaves whereas Taiwania has scale-like leaves. Both these genera have more extensive distributions in the fossil record, with close relatives of both known from Europe and North America.

Characters (from Watson & Eckenwalder): Trees or shrubs, evergreen (rarely deciduous), generally resinous and aromatic, monoecious (sometimes dioecious). Bark fibrous and furrowed (occasionally smooth or exfoliating in plates). Lateral branches well developed, similar to leading shoots, twigs terete, angled, or flattened dorsiventrally (with structurally distinct lower and upper surfaces), densely clothed by scalelike leaves or by decurrent leaf bases; longest internodes to 1 cm; buds usually undifferentiated and inconspicuous. Roots fibrous to woody (rarely bearing aboveground "knees"). Leaves simple, usually persisting 3-5 years and shed with lateral shoots (cladoptosic) (rarely shed annually), alternate and spirally arranged but sometimes twisted so as to appear 2-ranked, or opposite in 4 ranks, or whorled, deltate-scalelike to linear, decurrent, sessile or petioled; adult leaves appressed or spreading, often differing between lateral and leading shoots (twigs heterophyllous), sometimes strongly dimorphic on each twig with lateral scale-leaf pairs conspicuously keeled; juvenile leaves linear, flattened, spreading; often with solitary abaxial resin gland; resin canal present. Pollen cones maturing and shed annually, solitary, terminal (rarely in clusters of 2-5, axillary, or in terminal panicles), simple, spheric to oblong; sporophylls overlapping, bearing 2-10 abaxial microsporangia (pollen sacs); pollen spheric, not winged. Seed cones maturing in 1-2 seasons, shed with short shoots or persisting indefinitely on long-lived axes (rarely shattering at maturity), compound, solitary, terminal (rarely in clusters of 2-5 or axillary); scales overlapping or abutting, fused to subtending bracts with only bract apex sometimes free; each scale-bract complex peltate, oblong or cuneate, at maturity woody or fleshy, with 1-20 erect (rarely inverted with age), adaxial ovules. Seeds 1-20 per scale, not winged or with 2-3 symmetric or asymmetric wings; aril lacking; cotyledons 2-9.

Cupressaceae [Taxodiaceae, Taxodinae, Thujopsidae]
    |--Sewardiodendron laxum MM12
    |--Austrohamia MM12
    |    |--*A. minuta MM12
    |    `--A. acanthobractea MM12
    |--+--Cunninghamiostrobus yubariensis MM12
    |  |--Cunninghamites MM12
    |  |    |--C. elegans GT02
    |  |    `--C. lignitum MM12
    |  `--Cunninghamia [Cunninghamioideae] MM12
    |       |--C. hornbyensis MM12
    |       |--C. konishii MM12
    |       `--C. lanceolata MM12
    `--+--Taiwania [Taiwanioideae] MM12
       |    |--T. cretacea MM12
       |    |--T. cryptomerioides K02
       |    |--T. flousiana GT02
       |    `--T. schaeferi MM12
       `--+--Athrotaxidoideae MM12
          |    |--Athrotaxites berryi MM12
          |    `--Athrotaxis MM12
          |         |  i. s.: A. ungeri MM12
          |         |--A. cupressoides MM12
          |         `--+--A. laxifolia MM12
          |            `--A. selaginoides MM12
          `--+--+--Taxodioideae MM12
             |  `--+--Callitroideae MM12
             |     `--Cupressoideae MM12
             `--Sequoioideae MM12
                  |--Metasequoia MM12
                  |    |--M. foxii MM12
                  |    |--M. glyptostroboides MM12
                  |    |--M. milleri MM12
                  |    `--M. occidentalis MM12
                  `--+--Austrosequoia wintonensis MM12
                     |--Quasisequoia MM12
                     |    |--Q. couttsiae (Heer) Kunzmann 1999 (see below for synonymy) HL08
                     |    `--Q. florinii MM12
                     `--+--Sequoiadendron giganteum MM12
                        `--Sequoia MM12
                             |--S. ambigua GT02
                             |--S. carbonaria JD05
                             |--S. concinna GT02
                             |--S. gigantea H04
                             |--S. langsdorfi S89
                             |--S. reichenbachii MM12
                             `--S. sempervirens GT02

Cupressaceae incertae sedis:
  Geinitzia reichenbachii GT02
  Taxodioxylon gypsaceum JD05
  Cupressoconus rhenanus JD05
  Otakauia Pole 1995 P00
    |--*O. lanceolata P00
    `--O. macchatiensis Pole 2000 P00
  Cupressinoxylon (nom. cons.) JD05 [=Retinodendron (nom. rej.) S03]
    |--C. glyptostrobium JD05
    |--C. polonicum JD05
    |--C. ponderosum JD05
    `--C. sequoianum JD05
  Hughmillerites juddii MM12
  Elatides williamsonii MM12

Quasisequoia couttsiae (Heer) Kunzmann 1999 [=Sequoia couttsiae Heer 1862, Athrotaxis couttsiae (Heer) Gardner 1884] HL08

*Type species of generic name indicated

REFERENCES

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[HL08] Henniger, M., & R. Leder. 2008. Eozäne Makroflora der Querfurter Mulde. Mauritiana 20 (2): 229–251.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185–273.

[K02] Kvaček, Z. 2002. Novelties on Doliostrobus (Doliostrobaceae), an extinct conifer genus of the European Palaeogene. Časopis Národního Muzea, Řada Přírodovědná 171 (1–4): 47–62.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

[S03] Skog, J. E. 2003. Report of the Committee for Fossil Plants: 4. Taxon 52: 341.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 73–76.

Spermatophyta

Reconstruction of male reproductive structure of Sahnia nipaensis, from here.

Belongs within: Apo-Spermatophyta.
Contains: Cycadophyta, Coniferophyta, Arberiaceae, Pan-Gnetophyta, Pan-Angiospermae.

The name Spermatophyta was defined by Cantino et al. (2007) to represent the crown group of seed-bearing plants. Relationships within the spermatophytes remain uncertain: most morphological studies have supported a nested position for angiosperms among the remaining living taxa, but molecular analyses have supported a sister relationship between the angiosperms and living gymnosperms.

The clade Pan-Cycadophyta includes the cycads and all fossil taxa more closely related to them than other living seed plants. Known members of this clade are characterised by girdling leaf trances (Cantino et al. 2007), where leaf traces or veins leading to the leaf arise opposite the leaf attachment and circle the trunk within the storage tissue (Cycad Pages).

Fossil spermatophytes outside the major living lineages (cycads, conifers, gnetaleans and angiosperms) are mostly of the pteridosperm ('seed fern') grade, but some such as members of the Permian genus Glossopteris were large trees with entire leaves. The Pentoxylaceae are a group of plants (probably shrubs and small trees) known from the Jurassic, characterised by polystelic stems with a separate arrangement of vascular tissues for each stele. The Emplectopteridaceae were Permian pteridosperms with pinnate or entire leaves bearing flexuous or loosely anastomosed venation, and bilateral ovules attached to the underside of the leaves.

Possible synapomorphies (from Cantino et al. 2007): Endarch primary xylem in the stem, meiospores and microgametophytes with distal aperture, linear tetrad of megaspores, platyspermic ovules.

Spermatophyta [Acrogymnospermae, Phanerogamia, Pinophyta, Spermatophytatinae, Spermatophytina, Spermatopsida]
    |--Pan-Cycadophyta [Cycadopsida] CD07
    |    |  i. s.: Anomozamites RY03
    |    |         Antarcticycas CD07
    |    |         Michelilloa CD07
    |    |--Nilssoniaceae [Nilssoniales] Z02
    |    `--+--Semionogyna Z02
    |       `--Cycadophyta FC07
    `--+--Coniferophyta FC07
       `--+--Arberiaceae FC07
          |--+--Pan-Gnetophyta FC07
          |  `--Pan-Angiospermae FC07
          `--Pentoxylaceae [Pentoxylales] RL09
               |  i. s.: ‘Taeniopteris’ spatulata McClelland 1850 C93
               |--Pentoxylon sahnii Srivastava 1944 FC07, C93
               |--Nympaniophyllum hobsoni RL09
               |--Nipanioxylon guptai Srivastava 1944 C93
               |--Nipaniophyllum raoi Sahni 1948 C93
               |--Carnoconites RL09
               |    |--C. compactum Srivastava 1944 C93
               |    `--C. rajmahalensis (Wieland) Bose et al. 1984 C93
               `--Sahnia RL09
                    |--S. laxiphora PL12
                    `--S. nipaniensis Vishnu-Mittre 1953 C93

Spermatophyta incertae sedis:
  Emplectopteridaceae [Gigantonomiales, Gigantopteridaceae] Z02
    |--Gigantotheca paradoxa Li & Yao 1983 C93
    |--Emplectopteris Halle 1927 C93
    |    `--E. triangularis Halle 1927 C93
    |--Gigantonomia Li & Yao 1983 C93
    |    `--G. fukiensis (Yabe & Oishi) Li & Yao 1983 C93
    `--Gigantopteris Schenk 1883 C93
         |--G. americana D56
         `--G. taiyuanensis (Asama) Meyen 1987 C93
  Sanmiguelia PL12
  Balmeiopsis limbatus (Balme) Archangelsky 1979 I02
  Elaterocolpites castelainii Jardiné & Magloire 1965 I02
  Elateroplicites africaensis Herngreen 1973 I02
  Elaterosporites I02
    |--E. klaszii (Jardiné & Magloire) Jardiné 1967 I02
    `--E. verrucatus I02
  Galeacornea I02
    |--G. causea Stover 1963 I02
    `--G. clavis Stover 1963 I02
  Steevesipollenites sinuosus Azema & Boltenhagen 1974 I02
  Herodontia mikanioides J87
  Scolosanthus J87
    |--S. densiflorus J87
    `--S. selleanus J87
  Graffenrieda chrysandra J87
  Hottea torbeciana J87
  Reedia spathacea JK08
  Cycadopites PL12
    |--C. carpentieri (Delcourt & Sprumont) Singh 1964 I02
    |--C. durhamensis PL12
    |--C. follicularis PL12
    `--C. fragilis Singh 1964 I02
  Scalesia pedunculata HSS13
  Synangispadixis tidwellii PL12
  Cacabus miersii RJ11
  Leucosphaera Gilg 1897 KC01
  Monoceras Gothan 1909 KC01
  Preussiella Gilg 1897 KC01
  Rotularia Sternb. 1825 KC01

*Type species of generic name indicated

REFERENCES

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1–E44.

[C93] Cleal, C. J. 1993. Gymnospermophyta. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 795–808. Chapman & Hall: London.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549–552.

[HSS13] Hirschfeld, E., A. Swash & R. Still. 2013. The World's Rarest Birds. Princeton University Press: Princeton (New Jersey).

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775–788.

[JK08] Johnstone, R. E., & T. Kirkby. 2008. Distribution, status, social organisation, movements and conservation of Baudin's cockatoo (Calyptorhynchus baudinii) in south-west Western Australia. Records of the Western Australian Museum 25 (1): 107–118.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby's Dictionary of the Fungi 9^th ed. CAB International: Wallingford (UK).

[PL12] Peñalver, E., C. C. Labandeira, E. Barrón, X. Delclòs, P. Nel, A. Nel, P. Tafforeau & C. Soriano. 2012. Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the USA 109 (22): 8623–8628.

[RL09] Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C. Shih, A. Bashkuev, M. A. V. Logan, C. L. Hotton & D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326: 840–847.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1–11.

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

Last updated: 31 January 2020.