Revisions to the fossil sporophyte record of Marsilea
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Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U.S.A.
Department of Environmental and Plant Biology, Ohio University, Porter Hall 315, Athens, Ohio 45701, U.S.A.
Online publication date: 2019-06-18
Publication date: 2019-06-18
Acta Palaeobotanica 2019; 59(1): 27–50
The fossil record of Marsilea is challenging to assess, due in part to unreliable reports and conflicting opinions regarding the proper application of the names Marsilea and Marsileaceaephyllum to fossil leaves and leaflets similar to those of modern Marsilea. Specimens examined for this study include material assigned to Marsileaceaephyllum johnhallii, purportedly the oldest fossil record of a Marsilea-like sporophyte from the Lower Cretaceous of the Dakota Formation, Kansas, U.S.A.; leaves and leaf whorls of the extinct aquatic angiosperm Fortuna from several Late Cretaceous and Paleocene localities in western North America; and leaves and leaflets resembling Marsilea from the Eocene Green River Formation, Colorado and Utah, U.S.A. Literature on the fossil record of Marsilea was also reviewed. As a result, several taxonomic changes are proposed. Marsileaceaephyllum johnhallii is reinterpreted as an aquatic angiosperm that shares some architectural features with the genus Fortuna, although Marsileaceaephyllum is here maintained as a distinct genus with an emended diagnosis; under this reinterpretation, the name Marsileaceaephyllum can no longer be applied to sporophyte organs with affinities to Marsileaceae. Three valid fossil Marsilea species are recognized on the basis of sporophyte material that includes characteristic quadrifoliolate leaves and reticulate-veined leaflets: Marsilea campanica (J. Kvaček & Herman) Hermsen, comb. nov., from the Upper Cretaceous Grünbach Formation, Austria; Marsilea mascogos Estrada-Ruiz et al., from the Upper Cretaceous Olmos Formation, Mexico; and Marsilea sprungerorum Hermsen, sp. nov., from the Eocene Green River Formation, U.S.A. The species are distinguished from one another based on leaflet dimensions. Leaves from the Eocene Wasatch Formation, U.S.A., are transferred from Marsileaceaephyllum back to Marsilea, although not assigned to a fossil species. Finally, an occurrence of Marsilea from the Oligocene of Ethiopia is reassigned to Salvinia. A critical evaluation of the fossil record of Marsilea thus indicates that (1) the oldest fossil marsileaceous sporophytes bearing Marsilea-like leaves are from the Campanian; (2) only four credible records of sporophyte material attributable to Marsilea are known; and (3) the oldest dispersed Marsilea spores are known from the Oligocene.
Allen S.E. 2015. Fossil palm flowers from the Eocene of the Rocky Mountain region with affinities to Phoenix L. (Arecaceae: Coryphoideae). Int. J. Pl. Sci., 176(6): 586–596.
Aulenback K.R. 2009. Identification guide to the fossil plants of the Horseshoe Canyon Formation of Drumheller, Alberta. University of Calgary Press, Calgary.
Banerji J. 1987. Further contribution to the Mesozoic flora of Kutch, Gujarat. Geophytology, 17(1): 69–74.
Banerji J. 2004. Mesozoic megaflora of Kachchh Basin and its palaeoecological interpretation: 199–206. In: Srivastava P.C., (ed.), Vistas in palaeobotany and plant morphology: Evolutionary and environmental perspectives (Professor D.D. Pant memorial volume). U.P. Offset, Lucknow.
Batten D.J. & Kovach w.l. 1990. Catalog of Meso­zoic and Tertiary megaspores. Contr. Ser. Amer. Assoc. Stratigr. Palynologists, 24: 1–227.
Batten D.J., Collinson m.e. & Brain a.p.r. 2011. Megaspores and microspores of the extant and Paleogene marsileaceous fern Regnellidium and Cretaceous Molaspora: Evolutionary and phytogeographic implications. Int. J. Pl. Sci., 172(9): 1087–1100.
Bell W.A. 1949. Uppermost Cretaceous and Paleocene floras of western Alberta. Bull. Geol. Surv. Canada, 13: 1–231.
Berry E.W. 1915. The age of the Cretaceous flora of southern New York and New England. J. Geol., 23(7): 608–618.
Boucher L.D., Manchester s.r. & Judd w.s. 2003. An extinct genus of Salicaceae based on twigs with attached flowers, fruits, and foliage from the Eocene Green River Formation of Utah and Colorado, USA. Amer. J. Bot., 90(9): 1389–1399.
Bradley W.H. 1931. Origin and microfossils of the oil shale of the Green River Formation of Colorado and Utah. Profess. Pap. U.S. Geol. Surv., 168: 1–58.
Braun A. 1872. Ueber Marsilia Marioni, eine fossile Art aus der Tertiärzeit. Bot. Zeitung, 30(36): 653–654.
Brown R.W. 1934. The recognizable species of the Green River flora. Profess. Pap. U.S. Geol. Surv., 185–C: 45–77.
Call V.B. & Dilcher d.L. 1997. The fossil record of Eucommia (Eucommiaceae) in North America. Amer. J. Bot., 84(6): 798–814.
Chitaley S.D. & Paradkar s.a. 1972. Rodeites Sahni reinvestigated—I. Bot. J. Linn. Soc., 65(1): 109–117.
Christenhusz M.J.M. & Byng j.w. 2016. The number of known plants species in the world and its annual increase. Phytotaxa, 261(3): 201–217.
Collinson M.E. 1991. Diversification of modern heterosporous pteridophytes: 119–150. In: Blackmore S. & Barnes S.H. (eds.), Pollen and spores. Systematics Association Special Volume 44. Clarendon Press, Oxford.
Collinson M.E. 1996. “What use are fossil ferns?”- 20 years on: with a review of the fossil history of extant pteridophyte families and genera: 349–394. In: Camus J.M., Gibby M., Johns R.J. (eds.), Pteridology in perspective. Royal Botanic Gardens, Kew.
Collinson M.E. 2001. Cainozoic ferns and their distribution. Brittonia, 53(2): 173–235.
Collinson M.E., Smith s.y., van Konijnenburg-van Cittert j.h.a., Batten d.j., van der Burgh j., Barke j. & Marone f. 2013. New observations and synthesis of Paleogene heterosporous water ferns. Int. J. Pl. Sci., 174(3): 350–363.
Crouch N.R., Klopper R.R., Burrows J.E., Burrows S.M. 2011. Ferns of southern Africa, a comprehensive guide. Struik Nature, Cape Town, South Africa.
Cúneo N.R., Hermsen e.j. & Gandolfo m.a. 2013. Regnellidium (Salviniales, Marsileaceae) macrofossils and associated spores from the Late Cretaceous of South America. Int. J. Pl. Sci., 174(3): 340–349.
Cúneo N.R., Gandolfo m.a., Zamaloa m.c. & Hermsen e. 2014. Late Cretaceous aquatic plant world in Patagonia, Argentina. PLoS One, 9(8): e104749. DOI:10.1371/journal.pone.0104749.
Cushman R.A. Jr. 1983. Palynology and paleoecology of the Fossil Butte Member of the Eocene Green River Formation in Fossil Basin, Lincoln County, Wyoming. Thesis, Master of Science. Loma Linda University, Loma Linda, California.
Czaja A. 2003. Paläokarpologische Untersuchungen von Taphozönosen des Unter- und Mittelmiozäns aus dem Braunkohlentagebau Berzdorf/Oberlausitz (Sachsen). Palaeontographica, B, Paläophytol., 265(1–6): 1–148.
Dillhoff R.M., Dillhoff t.a., Dunn r.e., Myers j.a. & Strömberg c.a.e. 2009. Cenozoic paleobotany of the John Day Basin, central Oregon: 135–164. In: O’Connor J.E., Dorsey R.J. & Madin I.P. (eds.), Volcanoes to vineyards: Geologic field trips through the dynamic landscape of the Pacific Northwest. Geological Society of America Field Guide 15. Geological Society of America.
Dorofeev P.I. 1951. Meotychni roslyny z okolyts’ Odesy. Bot. Zhurn. (Kiev), 8(3): 31–40.
Dorofeev P.I. 1955a. Meoticheskaya flora iz okrestnostey g. Odessy. Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 1, Fl. Sist. Vyssh. Rast., 11: 109–143.
Dorofeev P.I. 1955b. Sarmatskie rasteniya s rek Tiligula i Ju. Buga. Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 1, Fl. Sist. Vyssh. Rast., 11: 144–160.
Dorofeev P.I. 1959. Materialy k poznaniju miotsenovoy flory Rostovskoy oblasti (summary: Contribution to the study of Miocene floras of the Rostov region). Probl. Bot., 4: 143–189.
Dorofeev P.I. 1963a. Tretichnye flory Zapadnoy Sibiri (summary: The Tertiary floras of western Siberia). Akademiya Nauk S.S.S.R. English translation of pgs. 62–80, 88–90 by MHW, Department of the Secretary of State, Translation Bureau, Multilingual Services Division, Canada (1975).
Dorofeev P.I. 1963b. Tretichnye rastenija Kazakhstana (summary: Tertiary plants of Kazakhstan). Bot. Zhurn. (Moscow & Leningrad), 48(2): 171–181.
Dorofeev P.I. 1981. K sistematike tretichnykh Marsileaceae (summary: On the taxonomy of the Tertiary Marsileaceae). Bot. Zhurn. (Moscow & Leningrad), 66(6): 792–801.
Dorofeev P.I. 1988. Miotsenovye flory Tambovskoy oblasti. Akademiya Nauk S.S.S.R. Botanischeskiy Institut im. V.L. Komarov, Leningrad.
Doweld A.B. 2016. Marsilea owambo, a new name for Marsilea vera (Marsileaceae). Amer. Fern J., 106(2): 143.
Ellis B., Daly d.c., Hickey l.j., Johnson k.r., Mitchell j.d., Wilf p. & Wing s.l. 2009. Manual of leaf architecture. Cornell University Press, Ithaca, New York.
Estrada-Ruiz E., Centeno-González n.k., Aguilar-Arellano f. & Martínez-Cabre­ra H.I. 2018. New record of the aquatic fern Marsilea, from the Olmos Formation (Upper Campanian), Coahuila, Mexico. Int. J. Pl. Sci., 179(6): 487–496.
Fikácek M., Prokin a. & Angus r.b. 2011. A long-living species of the hydrophiloid beetles: Helophorus sibiricus from the early Miocene deposits of Kartashevo (Siberia, Russia). ZooKeys, 130: 239–254.
Florin R. 1919. Eine Übersicht der fossilen Salvinia-Arten mit besonderer Berücksichtigung eines Fundes von Salvinia formosa Heer im Tertiär Japans. Bulletin of the Geological Institution of the University of Upsala, 16: 243–260.
Forno I.W. 1983. Native distribution of the Salvinia auriculata complex and keys to species identification. Aquatic Bot., 17(1): 71–83.
Foster A.S. & Gifford E.m. Jr 1974. Comparative morphology of vascular plants, 2nd ed. W.H. Freeman and Company, San Francisco.
Frič A. & Bayer e. 1900. Studien im Gebiete der Böhmischen Kreideformation. Palaeontologische Untersuchungen der einzelnen Schichten. Perucer Schichten. Archiv der naturwissenschaftlichen Landesdurchforschung von Böhmen, 11(2): 1–180.
Gandolfo M.A., Cúneo n.r. & Hermsen e.j. 2014. Reporte preliminar sobre la paleoflora de la Formación La Colonia (Campaniano-Maastrichtiano, Cretácico tardío), Chubut, Patagonia, Argentina. Bol. Soc. Geol. Mex., 66(1): 11–23.
García Massini J.L. & Jacobs B.F. 2011. The effects of volcanism on Oligocene-age plant communities from the Ethiopian Plateau, and implications for vegetational resilience in a heterogeneous landscape. Rev. Palaeobot. Palynol., 164: 211–222.
García Massini J.L., Jacobs b.f. & Tabor n.j. 2010. Paleobotany and sedimentology of late Oligocene terrestrial strata from the northwestern Ethiopian Plateau. Palaeontologia Electronica, 13.1.6A: 51 pages.
Gaudet J.J. 1964. Morphology of Marsilea vestita. II. Morphology of the adult land and submerged leaves. Amer. J. Bot., 51(6): 591–597.
Gemmill C.E.C. & Johnson k.r. 1997. Paleoecology of a Late Paleocene (Tiffanian) megaflora from the Northern Great Divide Basin, Wyoming. Palaios, 12(5): 439–448.
Gnibidenko Z.N. & Semakov n.n. 2009. Paleomagnetism of boundary Oligocene-Miocene deposits in the Kompasskii Bor tract on the Tym River (Western Siberia). Izvestiya, Physics of the Solid Earth, 45(1): 70–79.
Golovneva L.B. 1991. Novyi rod Palaeotrapa (Trapaceae?) i novye vidy Quereuxia iz rarytkinskoy svity (Koriakskoe nagor’ye, maastrikht-daniy) (The new genus Palaeotrapa (Trapaceae?) and new species Quereuxia from the Rarytkin Series (the Koryak Upland, the Maastrichtian-Danian). Bot. Zhurn., 76(4): 601–609.
Grande L. 2013. The lost world of Fossil Lake: Snapshots from deep time. University of Chicago Press, Chicago.
Gupta K.M. 1962. Marsilea. Bot. Monogr. Council Sci. Industr. Res., India, 2: 1–109.
Hattingh E.R. 1961. Problem of Salvinia auriculata Aubl. and associated aquatic weeds on Kariba Lake. Weed Res., 1(4): 303–306.
Heer O. 1874. Die Kreide-Flora der Arctischen Zone, gegründet auf die von den schwedischen Expeditionen von 1870 und 1872 in Grönland und Spitzbergen gesammelten Pflanzen. Kongl. Svenska Vetensk. Acad. Handl., 12(6): 1–138.
Heer O. 1882. Flora fossilis Grölandica, Die fossile flora Grönlands. Flora fossilis arctica, Die Fossile Flora der Polarländer, vol. 6, no. 2. J. Wurster & Comp., Zürich.
Herman A.B. & Kvaček j. 2007. Early Campanian Grünbach flora of Austria: systematic composition and palaeoclimatic interpretations. Acta Palaeobot., 47(1): 37–55.
Herman A.B. & Kvaček j. 2010. Late Cretaceous Grünbach Flora of Austria. Naturhistorisches Museum Wien, Wien.
Hermsen E.J., Gandolfo m.a. & Cúneo n.r. 2014. New marsileaceous fossils from the Late Cretaceous of South America and a reevaluation of Marsileaceaephyllum. Pl. Syst. Evol., 300(2): 369–386.
Hoffman G.L. 2002. Paleobotany and paleoecology of the Joffre Bridge Roadcut locality (Paleocene), Red Deer, Alberta, 2nd ed. Master of Science Thesis. University of Alberta, Edmonton, Alberta, Canada.
Hoffman G.L. & Stockey R.A. 1999. Geological setting and paleobotany of the Joffre Bridge Roadcut fossil locality (Late Paleocene), Red Deer Valley, Alberta. Canad. J. Earth Sci., 36(12): 2073–2084.
Hollick A. 1894. Fossil salvinias, including description of a new species. Bull. Torrey Bot. Club, 21(6): 253–257, plate 205.
Hollick A. 1904. Additions to the palaeobotany of the Cretaceous formation on Long Island. No. II. Bull. New York Bot. Gard., 3(11): 403–418, plates 71–79.
Hollick A. 1906. The Cretaceous flora of southern New York and New England. Monographs of the United States Geological Survey 50. Government Printing Office, Washington.
Hu S. & Taylor D.W. 2014. Floristics and Paleoecology of an Early Cretaceous Flora from Jordan. Bull. Peabody Mus. Nat. Hist., 55(2): 153–170.
Hu S., Taylor D.W., Brenner G.J. & Basha S.H. 2008. A new marsilealean fern species from the Early Cretaceous of Jordan. Palaeoworld, 17(3–4): 235–245.
IFPNI International Editorial Board. 2014–2018. The International Fossil Plant Names Index.
Jarmolenko A.V. 1935. Verkhne-melovaya flora severo-zapadnogo Kara-tau (summary: The Upper Cretaceous flora of the north-western Kara-tau). Trudy Sredne-Aziatsk. Gosud. Univ., Ser. 8b, Bot., 28: 1–36.
Johnson D.M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Syst. Bot. Monogr., 11: 1–87.
Johnson D.M. 1993. 23. Marsileaceae Mirbel, water clover family: 331–335. In: Flora of North America Editorial Committee (ed.), Flora of North America north of Mexico. Vol. 2 (Pteridophytes and Gymnosperms). Oxford University Press, New York and Oxford. Accessed at:
Johnson K.R. & Plumb c. 1995. Common plant fossils from the Green River Formation at Douglas Pass, Colorado, and Bonanza, Utah: 121–130. In: Averett W.R. (ed.), The Green River Formation in Piceance Creek and Eastern Uinta Basins. Grand Junction Geological Society, Grand Junction, Colorado.
Jud N.A. & Hickey L.J. 2013. Potomacapnos apeleutheron gen. et sp. nov., a new Early Cretaceous angiosperm from the Potomac Group and its implications for the evolution of eudicot leaf architecture. Amer. J. Bot., 100(12): 2437–2449.
Julien M.H., Center T.D. & Tipping P.W. 2002. Floating fern (Salvinia): 17–32. In: Van Driesche R., Lyon S., Blossey B., Hoddle M., Reardon R. (eds.), Biological control of invasive plants in the eastern United States. USDA Forest Service Publication FHTET-2002-04. Forest Health Technology Enterprise Team, Morgantown, West ­Virginia.
Knowlton F.H. 1900. Flora of the Montana Formation. Bull. U.S. Geol. Surv., 163: 1–118.
Knowlton F.H. 1902. Fossil flora of the John Day Basin Oregon. Bull. U.S. Geol. Surv., 204: 1–153.
Kovach W.L. & Batten D.J. 1989. Worldwide stratigraphic occurrences of Mesozoic and Tertiary megaspores. Palynology, 13(1): 247–277.
Kovach W.L. & Batten D.J. 1993. Diversity changes in lycopsid and aquatic fern megaspores through geologic time. Paleobiology, 19(1): 28–42.
Kovach W.L. & Dilcher D.L. 1988. Megaspores and other dispersed plant remains from the Dakota Formation (Cenomanian) of Kansas, U.S.A. Palynology, 12: 89–119.
Kramer K.U. 1990. Marsileaceae: 180–183. In: Kramer K.U. & Green P.S. (eds.), The families and genera of vascular plants. I. Pteridophytes and gymnosperms. Springer-Verlag, Berlin, Heidelberg.
Krasser F. 1906. Über die fossile Kreideflora von Grünbach in Niederösterreich. Anz. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl., 43(3): 41–43.
Kvaček J. & Herman A. 2004. The Campanian Grünbach flora of Lower Austria: palaeoecological interpretations. Ann. Naturhist. Mus. Wien, 106A: 91–101.
Launert E. 1968. A monographic survey of the genus Marsilea Linnaeus. The species of Africa and Madagascar. Senckenberg. Biol., 49(3/4): 273–315.
Lebedev I.V. 1955. Karaldinskiy kompleks: 157–158. In: Khalfin L.L. (ed.), Atlas rukovodyashchikh form iskopaemykh fauny i flory Zapadnoy Sibiri, tom II. Gosudarstvennoe Nauchno-Tekhnicheskoe Izdatel’stvo Literatury po Geologii i Okhrane Nedr, Moscow.
Lesquereux L. 1876. On the Tertiary flora of the North American lignitic, considered as evidence of the age of the formation. Rep. (Annual) U.S. Geol. Geogr. Surv. Territ., (1874): 271–365, plates 1–8.
Lesquereux L. 1878. Contributions to the fossil flora of the Western Territories. Part II. The Tertiary flora. Report of the United States Geological Survey of the Territories, vol. 7. Government Printing Office, Washington.
Lesquereux L. 1888. Recent determinations of fossil plants from Kentucky, Louisiana, Oregon, California, Alaska, Greenland, etc., with descriptions of new species. Proc. U.S. Natl. Mus., 11: 11–38, plates 4–16.
Lin Y.X. & Johnson d.m. 2013. Marsileaceae: 123–124. In: Wu Z.Y., Raven P.H. & Hong D.Y. (eds), Flora of China. Vol. 2–3 (Pteridophytes). Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. Accessed at:
Lupia R., Schneider H., Moeser G.M., Pryer K.M. & Crane P.R. 2000. Marsileaceae sporocarps and spores from the Late Cretaceous of Georgia, U.S.A. Int. J. Pl. Sci., 161(6): 975–988.
MacGinitie H.D. 1969. The Eocene Green River flora of northwestern Colorado and northeastern Utah. Univ. Calif. Publ. Geol. Sci., 83: 1–203.
Mai D.H. 1999. Die untermiozänen Floren aus der Spremberger Folge und dem 2. Flözhorizont in der Lausitz. Teil I: Farnpflanzen, Koniferen und Monokotyledonen. Palaeontographica, B, Paläophytol., 250(1–3): 1–76.
Mai D.H. 2000. Die mittelmiozänen und obermiozänen Floren aus der Meuroer und Raunoer Folge in der Lausitz Teil I: Farnpflanzen, Koniferen und Monoktoyledonen. Palaeontographica, B, Paläophytol., 256(1–3): 1–68.
Mai D.H. 2004. Die miozänen und pliozänen Floren aus Nordostbrandenburg und Südwestmecklenburg. Palaeontographica, B, Paläophytol., 269(1–6): 1–130.
Manchester S.R. 1989. Attached reproductive and vegetative remains of the extinct American-European genus Cedrelospermum (Ulmaceae) from the Early Tertiary of Utah and Colorado. Amer. J. Bot., 76(2): 256–276.
Manchester S.R. & Kodrul T.M. 2014. Morphology, affinities and phytogeographic history of Porosia Hickey in the Cretaceous and Paleocene of North America and Asia. Acta Palaeobot., 54(1): 77–99.
Manchester S.R., Crane P.R. & Golovneva L.B. 1999. An extinct genus with affinities to extant Davidia and Camptotheca (Cornales) from the Paleocene of North America and eastern Asia. Int. J. Pl. Sci., 160(1): 188–207.
Manchester S.R., Judd W.S. & Handley B. 2006. Foliage and fruits of early poplars (Salicaceae: Populus) from the Eocene of Utah, Colorado, and Wyoming. Int. J. Pl. Sci., 167(4): 897–908.
Marion A.-F. 1872. Plantes fossiles des calcaires marneux de Ronzon. Ann. Sci. Nat., Bot., 14: 326–364, plates 22–23.
Martínez-Hernández E., Almeida-Leñero L., Reyes-Salas M. & Betancourt-Aguilar Y. 1980. Estudio palinologico para la determinacion de ambientes en la cuenca Fuentes-Rio Escondido (Cretacico Superior), region de Piedras Negras, Coahuila. Revista del Instituto de Geología UNAM, 4(2): 167–185.
Matsunaga K.K.S., Smith S.Y., Manchester S.R., Kapgate D., Ramteke D., Garbout A., Villarraga-Gómez H. 2018. Reinvestigating an enigmatic Late Cretaceous monocot: morphology, taxonomy, and biogeography of Viracarpon. PeerJ, 6: e4580. DOI:10.7717/peerj.4580.
McIver E.E. & Basinger j.f. 1993. Flora of the Ravenscrag Formation (Paleocene), southwestern Saskatchewan, Canada. Palaeontographica Canadiana No.10. Canadian Society of Petroleum Geologists and Geological Association of Canada.
Merriam J.C. 1901. A contribution to the Geology of the John Day Basin. Bull. Dept. Geol. Univ. Calif., 2(9): 269–314.
Mitchell D.S. & Tur N.M. 1975. The rate of growth of Salvinia molesta (S. auriculata auct.) in laboratory and natural conditions. J. Appl. Ecol., 12(1): 213–225.
Monje-Dussán C., Martínez C., Escapa I. & Madriñán S. 2016. Nuevos registros de helechos y coníferas del Cretácico Inferior en la cuenca del Valle Superior del Magdalena, Colombia. Boletín de Geología, 38(4): 29–42.
Nagalingum N.S. 2007. Marsileaceaephyllum, a new genus for marsileaceous macrofossils: Leaf remains from the Early Cretaceous (Albian) of southern Gondwana. Pl. Syst. Evol., 264(1–2): 41–55.
Nagalingum N.S., Schneider H. & Pryer K.M. 2006. Comparative morphology of reproductive structures in heterosporous water ferns and a reevaluation of the sporocarp. Int. J. Pl. Sci., 167(4): 805–815.
Nagalingum N.S., Schneider H. & Pryer K.M. 2007. Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus Marsilea. Syst. Bot., 32(1): 16–25.
Nambudiri E.M.V. & Chitaley S. 2004. A permineralized fern rhizome with roots from the Upper Cretaceous (Maastrichtian) Deccan Intertrappean beds of India: 179–187. In: Srivastava P.C. (ed.), Vistas in palaeobotany and plant morphology: Evolutionary and environmental perspectives (Professor D.D. Pant memorial volume). U.P. Offset, Lucknow.
Negru A.G. 1986. Meoticheskaja flora severo-zapadnogo Prichernomor’ja. Shtiintsa, Kishinev.
Newman K.R. 1974. Palynomorph zones in Early Tertiary formations of the Piceance Creek and Uinta Basins, Colorado and Utah: 47–55. In: Murray D.K. (ed.), Energy resources of the Piceance Creek Basin, Colorado. 25th Field Conference, Rocky Mountain Association of Geologists, Denver, Colorado.
Paradkar S.A. 1971. Rhizomites dakshini gen. et. sp. nov. A new pteridophytic axis from the Deccan Intertrappean Beds of India. Botanique, 2(1): 15–19.
Paradkar S.A. 1988. A review of Indian fossil pteridophytes from the Deccan Intertrappean Beds. Geological Survey of India Special Publication No. 11, Vol 2: 175–183.
Paradkar S.A. & Barlinge s.g. 1981. Rodeites reinvestigated – III. Geophytology, 11(1): 16–24, plate 1.
Patil S.P., Kapgate d.k. & Zilpe s.k. 2014. Report of a fossil Marsilea petiole from the Deccan Intertrappean Beds of Maraipatan, Chandrapur District (M.S.). Pteridol. Res., 3(2): 10–14.
Pérez-Consuegra N., Cuervo-Gómez a., Martínez c., Montes c., Herrera f., Madriñán s. & Jaramillo c. 2017. Paleogene Salvinia (Salviniaceae) from Colombia and their paleobiogeographic implications. Rev. Palaeobot. Palynol., 246: 85–108.
PPG I [The Pteriophyte Phylogeny Group]. 2016. A community-derived classification for extant lycophytes and ferns. J. Syst. Evol., 54(6): 563–603.
Pray T.R. 1962. Ontogeny of the closed dichotomous venation of Regnellidium. Amer. J. Bot., 49(5): 464–472.
Pryer K.M. & Hearn d.j. 2009. Evolution of leaf form in marsileaceaceous ferns: Evidence for heterochrony. Evolution, 63(2): 498–513.
Puebla G.G., Prámparo M.B. & Gandolfo M.A. 2015. Aquatic ferns from the Upper Cretaceous Loncoche Formation, Mendoza, central-western, Argentina. Pl. Syst. Evol., 301(2): 577–588.
Reed C.F. 1954. Index Marsileata et Salviniata. Bol. Soc. Brot., ser. 2, 28: 1–61.
Reed C.F. 1965. Index Marsileata et Salviniata supplement. Bol. Soc. Brot., ser. 2, 39: 259–302.
Rich F.J., Johnson D.M. & Durkin T.V. 2001. Occurrence and paleoecology of Marsilea from the Eocene Wasatch Formation, Johnson County, Wyoming. Palaios, 16(6): 608–613.
Ruggiero M.A., Gordon D.P., Orrell T.M., Bailly N., Bourgoin T., Brusca R.C., Cavalier-Smith T., Guiry M.D., Kirk P.M. 2015. A higher level classification of all living organisms. PLoS ONE, 10(4): e0119248.
Schneider H. & Pryer K.M. 2002. Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. Int. J. Pl. Sci., 163(4): 485–505.
Schneller J.J. 1990. Salviniaceae: 256–258. In: Kramer K.U. & Green P.S. (eds.), The families and genera of vascular plants. I. Pteridophytes and gymnosperms. Springer-Verlag, Berlin, Heidelberg.
Sender L.M., Villanueva-Amadoz U., Diez J.B., Fernández-Baldor F.T. & Ferrer J. 2014. Earliest record of semi-aquatic fern leaves (family Marsileaceae) in the Mesozoic of Eurasia (Lower Cretaceous, Spain). Cretaceous Research, 51: 241–247.
Seward A.C. 1910. Fossil plants, a text-book for students of botany and geology, vol. 2. University Press, Cambridge.
Seward A.C. 1927. The Cretaceous plant-bearing rocks of western Greenland. Philos. Trans., Ser. B, 215: 57–175.
Shirley J. 1902. Notes on fossil plants from Duaringa, Ipswich, Dawson River, and Stanwell. Queensland Department of Mines Geological Survey Bulletin, 18: 7–13.
Shorthouse D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps.
Skog J.E. & Dilcher D.L. 1992. A new species of Marsilea from the Dakota Formation in central Kansas. Amer. J. Bot., 79(9): 982–988.
Skog J.E. & Dilcher D.L. 1994. Lower vascular plants of the Dakota Formation in Kansas and Nebraska, USA. Rev. Palaeobot. Palynol., 80(1–2): 1–18.
Smith A.R., Pryer k.m., Schuettpelz e., Korall p., Schneider h. & Wolf p.g. 2006. A classification for extant ferns. Taxon, 55(3): 705–731.
Smith M.E. & Carroll a.r. 2015. Introduction to the Green River Formation: 1–12. In: Smith M.E. & Carroll A.R. (eds.), Stratigraphy and paleolimnology of the Green River Formation, western USA. Syntheses in Limnogeology, vol. 1. Springer Science+Business Media, Dordrecht.
Srivastava R. & Guleria J.S. 2006. A catalogue of Cenozoic (Tertiary) plant megafossils from India (1989–2005). Birbal Sahni Institute of Palaeobotany, Lucknow.
Stockey R.A. & Rothwell G.W. 1997. The aquatic angiosperm Trapago angulata from the Upper Cretaceous (Maastrichtian) St. Mary River Formation of southern Alberta. Int. J. Pl. Sci., 158(1): 83–94.
Stockey R.A., Rothwell G.W. & Johnson K.R. 2007. Cobbania corrugata gen. et comb. nov. (Araceae): A floating aquatic monocot from the Upper Cretaceous of western North America. Amer. J. Bot., 94(4): 609–624.
Sun C., Li T., Na Y., Wu W., Li Y., Wang L. & Zhang L. 2014. Flabellariopteris, a new aquatic fern leaf from the Late Triassic of western Liaoning, China. Chin. Sci. Bull., 59(20): 2410–2418.
Tryon A.F. & Lugardon B. 1991. Spores of the Pteridophyta. Springer-Verlag, New York.
Tryon R.M. & Tryon a.f. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.
Upadhye E.V. & Patil g.v. 1979. On Marsilea like fossil rhizome and petiole from Deccan Intertrappean beds of Mohgaonkalan, India. J. Indian Bot. Soc., 58: 63–69.
Vallati P., De Sosa Tomas a., Casal g. & Calo m. 2017. Salviniales from the Late Cretaceous of the Golfo San Jorge Basin. Cretaceous Research, 74: 45–55.
Velenovský J. 1888. Ueber einige neue Pflanzenformen der böhmischen Kreideformation. Sitzungsber. Königl. Böhm Ges. Wiss. Prag, Math.-Naturwiss. Cl., Jahrgang 1887: 590–598.
Velenovský J. 1889. Květena Českého cenumanu. Rozpravy třídy mathematicko-přírodovědecké, Královské České Společnosti Náuk, 7(3): 1–75.
Velichkevich F.Yu. & Zastawniak E. 2003. The Pliocene flora of Kholmech, south-eastern Belarus and its correlation with other Pliocene floras of Europe. Acta Palaeobot., 43(2): 137–259.
Velichkevich F.Yu. & Zastawniak E. 2007. The state of investigation of the upper Pliocene Dvorets flora (SE Belarus). Acta Palaeobot., 47(1): 261–273.
WANG H. & DILCHER D.L. 2018. Early Cretaceous angiosperm leaves from the Dakota Formation, Hoisington III locality, Kansas, USA. Palaeontologia Electronica, 21.3.34A: 1–49.
Ward L.F. 1885. Sketch of paleobotany. Government Printing Office, Washington.
Wilf P. 2000. Late Paleocene–early Eocene climate changes in southwestern Wyoming: Paleobotanical analysis. Bull. Geol. Soc. Amer., 112(2): 292–307.
Wodehouse R.P. 1933 Tertiary Pollen-II, The oil shales of the Eocene Green River Formation. Bull. Torrey Bot. Club, 60(7): 479–524.
Yamada T. & Kato m. 2002. Regnellites nagashimae gen. et sp. nov., the oldest macrofossil of Marsileaceae, from the Upper Jurassic to Lower Cretaceous of western Japan. Int. J. Pl. Sci., 163(5): 715–723.
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