Evolution and Classification of Ferns and Lycophytes

  • Emily B. SessaEmail author


The last several decades have seen a transformation in our understanding of the evolutionary history of ferns and lycophytes. Historically the plants belonging to these lineages were grouped together and referred to as “pteridophytes” or “ferns and fern allies.” These terms both describe what we now recognize to be non-monophyletic assemblages, and both reflect a lack of understanding of the relationships of ferns and lycophytes that was only dispelled in the late 1990s and early 2000s. Thanks to the emergence of molecular systematics and the efforts of researchers in the fern and lycophyte community, we now have well-sampled and highly resolved phylogenies that include all tracheophytes, including substantial numbers of ferns and lycophytes. These trees have generated numerous insights about the evolutionary history of all land plants and have clarified the relationships between major groups of tracheophytes and vastly improved our understanding of fern and lycophyte evolution. This chapter reviews the major lineages, diversification events, and transitions in fern and lycophyte evolution and summarizes our current understanding of fern and lycophyte classification based on a modern, community-derived classification system that was recently produced for all 11,916 extant species of ferns and lycophytes.


Classification Divergence Diversification Evolution Fern Lycophyte Phylogeny Systematics Taxonomy 


  1. Almeida TE, Hennequin S, Schneider H, Smith AR, Batista JAN, Ramalho AJ, Proite K, Salino A (2016) Towards a phylogenetic generic classification of Thelypteridaceae: additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. Mol Phylogenet Evol 94:688–700PubMedCrossRefGoogle Scholar
  2. Arrigo N, Therrien J, Anderson CL, Windham MD, Haufler CH, Barker MS (2013) A total evidence approach to understanding phylogenetic relationships and ecological diversity in Selaginella subg. Tetragonostachys. Am J Bot 100:1672–1682PubMedCrossRefGoogle Scholar
  3. Assis FC, Almeida TE, Russell SJ, Schneider H, Salino A (2016) Molecular phylogeny and recircumscription of the fern genus Pecluma (Polypodiaceae–Polypodiopsida). Phytotaxa 247:235CrossRefGoogle Scholar
  4. Bateman RM, DiMichele WA (1994) Heterospory: the most iterative key innovation in the evolutionary history of the plant kingdom. Biol Rev 69:345–417CrossRefGoogle Scholar
  5. Bauret L, Gaudeul M, Sundue MA, Parris BS, Ranker TA, Rakotondrainibe F, Hennequin S, Ranaivo J, Selosse M-A, Rouhan G (2017) Madagascar sheds new light on the molecular systematics and biogeography of grammitid ferns: new unexpected lineages and numerous long-distance dispersal events. Mol Phylogenet Evol 111:1–17PubMedCrossRefGoogle Scholar
  6. Beck JB, Windham MD, Yatskievych G, Pryer KM (2010) A diploids-first approach to species delimitation and interpreting polyploid evolution in the fern genus Astrolepis (Pteridaceae). Syst Bot 35:223–234CrossRefGoogle Scholar
  7. Brouwer P, Bräutigam A, Külahoglu C, Tazelaar AOE, Kurz S, Nierop KGJ, van der Werf A, Weber APM, Schluepmann H (2014) Azolla domestication towards a biobased economy? New Phytol 202:1069–1082PubMedCrossRefGoogle Scholar
  8. Cardelus CL, Colwell RK, Watkins JE (2006) Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. J Ecol 94:144–156CrossRefGoogle Scholar
  9. Cárdenas GG, Tuomisto H, Lehtonen S (2016) Newly discovered diversity in the tropical fern genus Metaxya based on morphology and molecular phylogenetic analyses. Kew Bull 71:5CrossRefGoogle Scholar
  10. Chao Y-S, Rouhan G, Amoroso VB, Chiou W-L (2014) Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae). Ann Bot 114:109–124PubMedPubMedCentralCrossRefGoogle Scholar
  11. Christenhusz MJM, Byng JW (2016) The number of known plants species in the world and its annual increase. Phytotaxa 261:201–217CrossRefGoogle Scholar
  12. Christenhusz MJM, Zhang XC, Schneider H (2011) A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19:7CrossRefGoogle Scholar
  13. Clarke JT, Warnock RCM, Donoghue PCJ (2011) Establishing a time-scale for plant evolution. New Phytol 192:266–301PubMedCrossRefGoogle Scholar
  14. Collinson ME (2001) Cainozoic ferns and their distribution. Brittonia 53:173–235CrossRefGoogle Scholar
  15. Delevoryas T, Hueber F, Banks H, Beck C, Eggert DA, Arnold CA, White RA, Tryon RM, Wagner WH (1964) The origin and evolution of ferns. Mem Torrey Bot Club 21:3–95Google Scholar
  16. Der JP, Thomson JA, Stratford JK, Wolf PG (2009) Global chloroplast phylogeny and biogeography of Bracken (Pteridium; Dennstaedtiaceae). Am J Bot 96:1041–1049PubMedCrossRefGoogle Scholar
  17. Des Marais DL, Smith AR, Britton DM, Pryer KM (2003) Phylogenetic relationships and evolution of extant horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F). Int J Plant Sci 164:737–751CrossRefGoogle Scholar
  18. Dubuisson JY, Hennequin S, Douzery EJP, Cranfill R, Smith AR, Pryer KM (2003) rbcL phylogeny of the fern genus Trichomanes (Hymenophyllaceae), with special reference to Neotropical taxa. Int J Plant Sci 164:753–761CrossRefGoogle Scholar
  19. Ebihara A, Dubuisson J-Y, Iwatsuki K, Hennequin S, Ito M (2006) A taxonomic revision of Hymenophyllaceae. Blumea – Biodivers Evol Biogeogr Plants 51:221–280CrossRefGoogle Scholar
  20. Ebihara A, Iwatsuki K, Ito M, Hennequin S, Dubuisson J-Y (2007) A global molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Bot J Linn Soc 155:1–27CrossRefGoogle Scholar
  21. Field AR, Bostock PD (2013) New and existing combinations in Palaeotropical Phlegmariurus (Lycopodiaceae) and lectotypification of the type species Phlegmariurus phlegmaria (L.) T.Sen & U.Sen. PhytoKeys 20:33–51CrossRefGoogle Scholar
  22. Field AR, Testo W, Bostock PD, Holtum JAM, Waycott M (2016) Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum. Mol Phylogenet Evol 94:635–657PubMedCrossRefGoogle Scholar
  23. de Gasper AL, Dittrich VADO, Smith AR, Salino A (2016) A classification for Blechnaceae (Polypodiales: Polypodiopsida): new genera, resurrected names, and combinations. Phytotaxa 275:191CrossRefGoogle Scholar
  24. de Gasper AL, Almeida TE, Dittrich VAO, Smith AR, Salino A (2017) Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus-level treatment. Cladistics Int J Willi Hennig Soc 33:429–446CrossRefGoogle Scholar
  25. Gastaldo RA (1987) Confirmation of Carboniferous clastic swamp communities. Nature 326:869–871CrossRefGoogle Scholar
  26. Gensel PG (1984) Plant life in the devonian. Praeger, New YorkGoogle Scholar
  27. Gerrienne P, Servais T, Vecoli M (2016) Plant evolution and terrestrialization during Palaeozoic times—The phylogenetic context. Rev Palaeobot Palynol 227:4–18CrossRefGoogle Scholar
  28. Grewe F, Guo W, Gubbels EA, Hansen AK, Mower JP (2013) Complete plastid genomes from Ophioglossum californicum, Psilotum nudum, and Equisetum hyemale reveal an ancestral land plant genome structure and resolve the position of Equisetales among monilophytes. BMC Evol Biol 13:8PubMedPubMedCentralCrossRefGoogle Scholar
  29. Grusz AL, Windham MD (2013) Toward a monophyletic Cheilanthes: the resurrection and recircumscription of Myriopteris (Pteridaceae). PhytoKeys:49–64Google Scholar
  30. Guillon J-M (2004) Phylogeny of horsetails (Equisetum) based on the chloroplast rps4 gene and adjacent noncoding sequences. Syst Bot 29:251–259CrossRefGoogle Scholar
  31. Guillon J-M (2007) Molecular phylogeny of horsetails (Equisetum) including chloroplast atpB sequences. J Plant Res 120:569–574PubMedCrossRefGoogle Scholar
  32. Hasebe M, Omori T, Nakazawa M, Sano T, Kato M, Iwatsuki K (1994) rbcL gene sequences provide evidence for the evolutionary lineages of leptosporangiate ferns. Proc Natl Acad Sci U S A 91:5730–5734PubMedPubMedCentralCrossRefGoogle Scholar
  33. Hasebe M, Wolf PG, Pryer KM, Ueda K, Ito M, Sano R, Gastony GJ, Yokoyama J, Manhart JR, Murakami N, Crane EH, Haufler CH, Hauk WD (1995) Fern phylogeny based on rbcL nucleotide sequences. Am Fern J 85:134CrossRefGoogle Scholar
  34. Haufler CH, Pryer KM, Schuettpelz E, Sessa EB, Farrar DR, Moran R, Schneller JJ, Watkins JE, Windham MD (2016) Sex and the single gametophyte: revising the homosporous vascular plant life cycle in light of contemporary research. Bioscience 66:928–937CrossRefGoogle Scholar
  35. He L-J, Zhang X-C (2012) Exploring generic delimitation within the fern family Thelypteridaceae. Mol Phylogenet Evol 65:757–764PubMedCrossRefGoogle Scholar
  36. Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson J-Y (2003) Molecular systematics of the fern genus Hymenophyllum s.l. (Hymenophyllaceae) based on chloroplastic coding and noncoding regions. Mol Phylogenet Evol 27:283–301PubMedCrossRefGoogle Scholar
  37. Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson JY (2006) Phylogenetic systematics and evolution of the genus Hymenophyllum (Hymenophyllaceae: Pteridophyta). Fern Gaz 17:247–257Google Scholar
  38. Hennequin S, Hovenkamp P, Christenhusz MJM, Schneider H (2010) Phylogenetics and biogeography of Nephrolepis – a tale of old settlers and young tramps. Bot J Linn Soc 164:113–127CrossRefGoogle Scholar
  39. Hickey RJ (1986) The early evolutionary and morphological diversity of Isoetes, with descriptions of two new neotropical species. Syst Bot 11:309CrossRefGoogle Scholar
  40. Holub J (1983) Validation of generic names in Lycopodiaceae: with a description of a new genus Pseudolycopodiella. Folia Geobotanica et Phytotaxonomica 18:439–442CrossRefGoogle Scholar
  41. Hoot SB, Taylor WC (2001) The utility of nuclear ITS, a LEAFY homolog intron, and chloroplast atpB-rbcL space region data in phylogenetic analyses and species delimitation in Isoetes. Am Fern J 91:166–177CrossRefGoogle Scholar
  42. Hoot SB, Taylor WC, Napier NS (2006) Phylogeny and biogeography of Isoëtes (Isoëtaceae) based on nuclear and chloroplast DNA sequence data. Syst Bot 31:449–460CrossRefGoogle Scholar
  43. Hueber FM (1992) Thoughts on the early lycopsids and zosterophylls. Ann Mo Bot Gard 79:474CrossRefGoogle Scholar
  44. Jermy AC (1986) Subgeneric names in Selaginella. Fern Gaz 13:117–118Google Scholar
  45. Ji SG, Huo KK, Wang J, Pan SL (2008) A molecular phylogenetic study of Huperziaceae based on chloroplast rbcL and psbA-trnH sequences. J Syst Evol 46:213–219Google Scholar
  46. Johnson DM (1986) Systematics of the new world species of Marsilea (Marsileaceae). Syst Bot Monogr 11:1CrossRefGoogle Scholar
  47. Kenrick P, Crane PR, Kenrick P, Crane P (1997) The origin and early evolution of plants on land. Nature 389:33–39CrossRefGoogle Scholar
  48. Kenrick P, Wellman CH, Schneider H, Edgecombe GD (2012) A timeline for terrestrialization: consequences for the carbon cycle in the Palaeozoic. Philos Trans R Soc Lond B Biol Sci 367:519–536PubMedPubMedCentralCrossRefGoogle Scholar
  49. Kirkpatrick REB (2007) Investigating the monophyly of Pellaea (Pteridaceae) in the context of a phylogenetic analysis of cheilanthoid ferns. Syst Bot 32:504–518CrossRefGoogle Scholar
  50. Klekowski EJ, Baker HG (1966) Evolutionary significance of polyploidy in the pteridophyta. Science 153:305–307PubMedCrossRefGoogle Scholar
  51. Knie N, Fischer S, Grewe F, Polsakiewicz M, Knoop V (2015) Horsetails are the sister group to all other monilophytes and Marattiales are sister to leptosporangiate ferns. Mol Phylogenet Evol 90:140–149PubMedCrossRefGoogle Scholar
  52. Korall P, Kenrick P (2002) Phylogenetic relationships in Selaginellaceae based on RBCL sequences. Am J Bot 89:506–517PubMedCrossRefGoogle Scholar
  53. Korall P, Kenrick P (2004) The phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: extreme substitution rates and rate heterogeneity. Mol Phylogenet Evol 31:852–864PubMedCrossRefGoogle Scholar
  54. Korall P, Pryer KM (2014) Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal. J Biogeogr 41:402–413PubMedCrossRefGoogle Scholar
  55. Korall P, Kenrick P, Therrien JP (1999) Phylogeny of selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. Int J Plant Sci 160:585–594CrossRefGoogle Scholar
  56. Korall P, Pryer KM, Metzgar JS, Schneider H, Conant DS (2006) Tree ferns: monophyletic groups and their relationships as revealed by four protein-coding plastid loci. Mol Phylogenet Evol 39:830–845PubMedCrossRefGoogle Scholar
  57. Korall P, Conant DS, Metzgar JS, Schneider H, Pryer KM (2007) A molecular phylogeny of scaly tree ferns (Cyatheaceae). Am J Bot 94:873–886PubMedCrossRefGoogle Scholar
  58. Kreier H-P, Schneider H (2006) Reinstatement of Loxogramme dictyopteris, based on phylogenetic evidence, for the New Zealand endemic fern, Anarthropteris lanceolata (Polypodiaceae, Polypodiidae). Aust Syst Bot 19:309CrossRefGoogle Scholar
  59. Kreier H-P, Zhang X-C, Muth H, Schneider H (2008) The microsoroid ferns: Inferring the relationships of a highly diverse lineage of Paleotropical epiphytic ferns (Polypodiaceae, Polypodiopsida). Mol Phylogenet Evol 48:1155–1167PubMedCrossRefGoogle Scholar
  60. Kumar K (2001) Reproductive biology of pteridophytes. In: Johri BM, Srivastava PS (eds) Reproductive biology of plants. Springer, New York, pp 175–214CrossRefGoogle Scholar
  61. Kuo L-Y, Li F-W, Chiou W-L, Wang C-N (2011) First insights into fern matK phylogeny. Mol Phylogenet Evol 59:556–566PubMedCrossRefGoogle Scholar
  62. Labiak PH, Sundue M, Rouhan G (2010) Molecular phylogeny, character evolution, and biogeography of the grammitid fern genus Lellingeria (Polypodiaceae). Am J Bot 97:1354–1364PubMedCrossRefGoogle Scholar
  63. Labiak PH, Sundue M, Rouhan G, Hanks JG, Mickel JT, Moran RC (2014) Phylogeny and historical biogeography of the lastreopsid ferns (Dryopteridaceae). Am J Bot 101:1207–1228PubMedCrossRefGoogle Scholar
  64. Labiak PH, Mickel JT, Hanks JG (2015a) Molecular phylogeny and character evolution of Anemiaceae (Schizaeales). Taxon 64:1141–1158CrossRefGoogle Scholar
  65. Labiak PH, Sundue M, Rouhan G, Moran RC (2015b) New combinations in Lastreopsis and Parapolystichum (Dryopteridaceae). Brittonia 67:79–86CrossRefGoogle Scholar
  66. Larsén E, Rydin C (2016) Disentangling the phylogeny of Isoetes (Isoetales), using nuclear and plastid data. Int J Plant Sci 177:157–174CrossRefGoogle Scholar
  67. Le Péchon T, He H, Zhang L, Zhou X-M, Gao X-F, Zhang L-B (2016a) Using a multilocus phylogeny to test morphology-based classifications of Polystichum (Dryopteridaceae), one of the largest fern genera. BMC Evol Biol 16:55PubMedPubMedCentralCrossRefGoogle Scholar
  68. Le Péchon T, Zhang L, He H, Zhou X-M, Bytebier B, Gao X-F, Zhang L-B (2016b) A well-sampled phylogenetic analysis of the polystichoid ferns (Dryopteridaceae) suggests a complex biogeographical history involving both boreotropical migrations and recent transoceanic dispersals. Mol Phylogenet Evol 98:324–336PubMedCrossRefGoogle Scholar
  69. Lehtonen S (2011) Towards resolving the complete fern tree of life. Plos One 6:e24851PubMedPubMedCentralCrossRefGoogle Scholar
  70. Lehtonen S, Tuomisto H, Rouhan G, Christenhusz MJM (2010) Phylogenetics and classification of the pantropical fern family Lindsaeaceae. Bot J Linn Soc 163:305–359CrossRefGoogle Scholar
  71. Lehtonen S, Wahlberg N, Christenhusz MJM (2012) Diversification of lindsaeoid ferns and phylogenetic uncertainty of early polypod relationships. Bot J Linn Soc Linn Soc Lond 170:489–503CrossRefGoogle Scholar
  72. Lenton TM, Dahl TW, Daines SJ, Mills BJW, Ozaki K, Saltzman MR, Porada P (2016) Earliest land plants created modern levels of atmospheric oxygen. Proc Natl Acad Sci U S A 113:9704–9709PubMedPubMedCentralCrossRefGoogle Scholar
  73. Li F-W, Pryer KM (2014) Crowdfunding the Azolla fern genome project: a grassroots approach. GigaScience 3:16PubMedPubMedCentralCrossRefGoogle Scholar
  74. Li C-X, Lu S-G, Barrington DS (2008) Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL-F and rps4-trnS). J Plant Res 121:19–26PubMedCrossRefGoogle Scholar
  75. Li C, Lu S, Sun X, Yang Q (2011) Phylogenetic positions of the enigmatic asiatic fern genera Diplaziopsis and Rhachidosorus from analyses of four plastid genes. Am Fern J 101:142–155CrossRefGoogle Scholar
  76. Li F-W, Pryer KM, Windham MD (2012) Gaga, a new fern genus segregated from Cheilanthes (Pteridaceae). Syst Bot 37:845–860CrossRefGoogle Scholar
  77. Link-Pérez MA, Watson LE, Hickey RJ (2011) Redefinition of Adiantopsis Fée (Pteridaceae): systematics, diversification, and biogeography. Taxon 60:1255–1268Google Scholar
  78. Linnaeus C (1753) Species plantarum. Laurentius Salvius, StockholmGoogle Scholar
  79. Little DP, Barrington DS (2003) Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). Am J Bot 90:508–514PubMedCrossRefGoogle Scholar
  80. Liu H-M, Jiang R-H, Guo J, Hovenkamp P, Perrie LR, Shepherd L, Hennequin S, Schneider H (2013) Towards a phylogenetic classification of the climbing fern genus Arthropteris. Taxon 62:688–700CrossRefGoogle Scholar
  81. Liu H-M, Zhang X-C, Wang M-P, Shang H, Zhou S-L, Yan Y-H, Wei X-P, Xu W-B, Schneider H (2016) Phylogenetic placement of the enigmatic fern genus Trichoneuron informs on the infra-familial relationship of Dryopteridaceae. Plant Syst Evol 302:319–332CrossRefGoogle Scholar
  82. Llorens C, Argentina M, Rojas N, Westbrook J, Dumais J, Noblin X (2016) The fern cavitation catapult: mechanism and design principles. J R Soc Interface 13:20150930PubMedPubMedCentralCrossRefGoogle Scholar
  83. Lóriga J, Vasco A, Regalado L, Heinrichs J, Moran RC (2014) Phylogeny and classification of the Cuban species of Elaphoglossum (Dryopteridaceae), with description of Elaphoglossum sect. Wrightiana sect. nov. Plant Syst Evol 300:937–951CrossRefGoogle Scholar
  84. Lovis JD (1977) Evolutionary patterns and processes in ferns. Adv Bot Res 4:229–440CrossRefGoogle Scholar
  85. Madeira PT, Pemberton RW, Center TD (2008) A molecular phylogeny of the genus Lygodium (Schizaeaceae) with special reference to the biological control and host range testing of Lygodium microphyllum. Biol Control 45:308–318CrossRefGoogle Scholar
  86. Magallón S, Hilu KW, Quandt D (2013) Land plant evolutionary timeline: gene effects are secondary to fossil constraints in relaxed clock estimation of age and substitution rates. Am J Bot 100:556–573PubMedCrossRefGoogle Scholar
  87. Manhart JR (1995) Chloroplast 16S rDNA sequences and phylogenetic relationships of fern allies and ferns. Am Fern J 85:182CrossRefGoogle Scholar
  88. Matos FB, Mickel JT (2014) The Brazilian species of Elaphoglossum section Polytrichia (Dryopteridaceae). Brittonia 66:371–395CrossRefGoogle Scholar
  89. Metzgar JS, Schneider H, Pryer KM (2007) Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae). Int J Plant Sci 168:1045–1053CrossRefGoogle Scholar
  90. Metzgar JS, Skog JE, Zimmer EA, Pryer KM (2008) The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Syst Bot 33:31–36CrossRefGoogle Scholar
  91. Metzgar JS, Alverson ER, Chen S, Vaganov AV, Ickert-Bond SM (2013) Diversification and reticulation in the circumboreal fern genus Cryptogramma. Mol Phylogenet Evol 67:589–599PubMedCrossRefGoogle Scholar
  92. Moran RC (2004) A natural history of ferns. Timber Press, Inc., PortlandGoogle Scholar
  93. Moran RC, Labiak PH (2015) Phylogeny of the polybotryoid fern clade (Dryopteridaceae). Int J Plant Sci 176:880–891CrossRefGoogle Scholar
  94. Moran RC, Labiak PH, Sundue M (2010) Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae). Int J Plant Sci 171:547–559CrossRefGoogle Scholar
  95. Mueller RJ (1982) Shoot morphology of the climbing fern Lygodium (Schizaeaceae): general organography, leaf initiation, and branching. Bot Gaz 143:319–330CrossRefGoogle Scholar
  96. Mueller RJ (1983) Indeterminate growth and ramification of the climbing leaves of Lygodium japonicum (Schizaeaceae). Am J Bot 70:682CrossRefGoogle Scholar
  97. Murdock AG (2008a) A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new genus Ptisana. Taxon 57:737–755Google Scholar
  98. Murdock AG (2008b) Phylogeny of marattioid ferns (Marattiaceae): inferring a root in the absence of a closely related outgroup. Am J Bot 95:626–641PubMedCrossRefGoogle Scholar
  99. Mynssen CM, Vasco A, Moran RC, Sylvestre LS, Rouhan G (2016) Desmophlebiaceae and Desmophlebium: a new family and genus of Eupolypod II ferns. Taxon 65:19–34CrossRefGoogle Scholar
  100. Nagalingum NS, Schneider H, Pryer KM (2007) Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus Marsilea. Syst Bot 32:16–25CrossRefGoogle Scholar
  101. Nagalingum NS, Nowak MD, Pryer KM (2008) Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia. Bot J Linn Soc 157:673–685CrossRefGoogle Scholar
  102. Nickrent DL, Parkinson CL, Palmer JD, Duff RJ (2000) Multigene phylogeny of land plants with special reference to bryophytes and the earliest land plants. Mol Biol Evol 17:1885–1895PubMedCrossRefGoogle Scholar
  103. Noblin X, Rojas NO, Westbrook J, Llorens C, Argentina M, Dumais J (2012) The fern sporangium: a unique catapult. Science 335:1322PubMedCrossRefGoogle Scholar
  104. Øllgaard B (2015) Six new species and some nomenclatural changes in neotropical Lycopodiaceae. Nord J Bot 33:186–196CrossRefGoogle Scholar
  105. Otto EM, Janssen T, Kreier H-P, Schneider H (2009) New insights into the phylogeny of Pleopeltis and related Neotropical genera (Polypodiaceae, Polypodiopsida). Mol Phylogenet Evol 53:190–201PubMedCrossRefGoogle Scholar
  106. Pabby A, Prasanna R, Singh PK (2004) Biological significance of Azolla and its utilization in agriculture. Proc Indian Natl Sci Acad B70:299–333Google Scholar
  107. Pemberton RW, Ferriter AP (1998) Old world climbing fern (Lygodium microphyllum), a dangerous invasive weed in Florida. Am Fern J 88:165CrossRefGoogle Scholar
  108. Pereira AL, Martins M, Oliveira MM, Carrapiço F (2011) Morphological and genetic diversity of the family Azollaceae inferred from vegetative characters and RAPD markers. Plant Syst Evol 297:213–226CrossRefGoogle Scholar
  109. Perkins SK, Peters GA (2006) The Azolla-Anabaena symbiosis: endophyte continuity in the Azolla life-cycle is facilitated by epidermal trichomes. New Phytol 123:53–64CrossRefGoogle Scholar
  110. Perrie LR, Shepherd LD, Brownsey PJ (2015) An expanded phylogeny of the Dennstaedtiaceae ferns: Oenotrichia falls within a non-monophyletic Dennstaedtia, and Saccoloma is polyphyletic. Aust Syst Bot 28:256CrossRefGoogle Scholar
  111. Phipps C, Taylor T, Taylor E, Cúneo R, Boucher L, Yao X (1998) Osmunda (Osmundaceae) from the Triassic of Antarctica: an example of evolutionary stasis. Am J Bot 85:888PubMedCrossRefGoogle Scholar
  112. Pittermann J, Watkins JE, Cary KL, Schuettpelz E, Brodersen C, Smith AR, Baer A (2015) The structure and function of xylem in seed-free vascular plants: an evolutionary perspective. In: Hacke U (ed) Functional and ecological xylem anatomy. Springer International Publishing, Cham, pp 1–37Google Scholar
  113. Plackett ARG, Coates JC (2016) Life’s a beach – the colonization of the terrestrial environment. New Phytol 212:831–835PubMedCrossRefGoogle Scholar
  114. PPG 1 (2016) A community-derived classification for extant lycophytes and ferns. J Syst Evol 54:563–603CrossRefGoogle Scholar
  115. Prado J, Del Nero Rodrigues C, Salatino A, Salatino ML (2007) Phylogenetic relationships among Pteridaceae, including Brazilian species, inferred from rbcL sequences. Taxon 56:355–368Google Scholar
  116. Pryer KM (1999) Phylogeny of marsileaceous ferns and relationships of the fossil Hydropteris pinnata reconsidered. Int J Plant Sci 160:931–954PubMedCrossRefGoogle Scholar
  117. Pryer KM, Smith AR, Skog JE, Present A-W (1995) Phylogenetic relationships of extant ferns based on evidence from morphology and rbcL sequences. Am Fern J 85:205–282CrossRefGoogle Scholar
  118. Pryer KM, Schneider H, Smith AR, Cranfill R, Wolf PG, Hunt JS, Sipes SD (2001a) Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Nature 409:618–622PubMedCrossRefGoogle Scholar
  119. Pryer KM, Smith AR, Hunt JS, Dubuisson JY (2001b) rbcL data reveal two monophyletic groups of filmy ferns (Hymenophyllaceae). Am J Bot 88:1118–1130PubMedCrossRefGoogle Scholar
  120. Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R (2004) Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. Am J Bot 91:1582–1598PubMedCrossRefGoogle Scholar
  121. Qiu YL, Yu J (2003) Azolla – a model organism for plant genomic studies. Genomics Proteomics Bioinformatics 1:15–25PubMedPubMedCentralCrossRefGoogle Scholar
  122. Qiu Y-L, Li L, Wang B, Chen Z, Knoop V, Groth-Malonek M, Dombrovska O, Lee J, Kent L, Rest J, Estabrook GF, Hendry TA, Taylor DW, Testa CM, Ambros M, Crandall-Stotler B, Duff RJ, Stech M, Frey W, Quandt D, Davis CC (2006) The deepest divergences in land plants inferred from phylogenomic evidence. Proc Natl Acad Sci U S A 103:15511–15516PubMedPubMedCentralCrossRefGoogle Scholar
  123. Qiu Y, Li L, Wang B, Chen Z, Dombrovska O, Lee J, Kent L, Li R, Jobson RW, Hendry TA, Taylor DW, Testa CM, Ambros M (2007) A nonflowering land plant phylogeny inferred from nucleotide sequences of seven chloroplast, mitochondrial, and nuclear genes. Int J Plant Sci 168:691–708CrossRefGoogle Scholar
  124. Rai HS, Graham SW (2010) Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes). Am J Bot 97:1444–1456PubMedCrossRefGoogle Scholar
  125. Ranker TA, Smith AR, Parris BS, Geiger JMO, Haufler CH, Straub SCK, Schneider H (2004) Phylogeny and evolution of grammitid ferns: a case of rampant morphological homoplasy. Taxon 53:415–428CrossRefGoogle Scholar
  126. Reid JD, Plunkett GM, Peters GA (2006) Phylogenetic relationships in the heterosporous fern genus Azolla (Azollaceae) based on DNA sequence data from three noncoding regions. Int J Plant Sci 167:529–538CrossRefGoogle Scholar
  127. Rothfels CJ, Schuettpelz E (2014) Accelerated rate of molecular evolution for vittarioid ferns is strong and not driven by selection. Syst Biol 63:31–54PubMedCrossRefGoogle Scholar
  128. Rothfels CJ, Larsson A, Kuo L-Y, Korall P, Chiou W-L, Pryer KM (2012a) Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Syst Biol 61:490–509PubMedCrossRefGoogle Scholar
  129. Rothfels CJ, Sundue MA, Kuo L-Y, Larsson A, Kato M, Schuettpelz E, Pryer KM, Rothfels C, Sundue M, Kuo L-Y, Larsson A, Kato M, Schuettpelz E, Pryer K (2012b) A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61:515–533Google Scholar
  130. Rothfels CJ, Windham MD, Pryer KM (2013) A plastid phylogeny of the cosmopolitan fern family cystopteridaceae (Polypodiopsida). Syst Bot 38:295–306CrossRefGoogle Scholar
  131. Rothfels CJ, Li F-W, Sigel EM, Huiet L, Larsson A, Burge DO, Ruhsam M, Deyholos M, Soltis DE, Stewart CN, Shaw SW, Pokorny L, Chen T, dePamphilis C, DeGironimo L, Chen L, Wei X, Sun X, Korall P, Stevenson DW, Pryer KM (2015) The evolutionary history of ferns inferred from 25 low-copy nuclear genes. Am J Bot 102:1089–1107PubMedCrossRefGoogle Scholar
  132. Rothwell GW (1987) Complex paleozoic filicales in the evolutionary radiation of ferns. Am J Bot 74:458CrossRefGoogle Scholar
  133. Rothwell GW (1999) Fossils and ferns in the resolution of land plant phylogeny. Bot Rev 65:188–218CrossRefGoogle Scholar
  134. Rothwell GW, Nixon KC (2006) How does the inclusion of fossil data change our conclusions about the phylogenetic history of euphyllophytes? Int J Plant Sci 167:737–749CrossRefGoogle Scholar
  135. Rothwell GW, Stockey RA (1994) The role of Hydropteris pinnata gen. et. sp. nov. in reconstructing the cladistics of heterosporous ferns. Am J Bot 81:479CrossRefGoogle Scholar
  136. Rothwell GW, Stockey RA (2006) Combining characters of pteridaceae and tree ferns: Pterisorus radiata gen. et sp. nov., a permineralized lower Cretaceous filicalean with radial sori. Int J Plant Sci 167:695–701CrossRefGoogle Scholar
  137. Rothwell GW, Stockey RA (2008) Phylogeny and evolution of ferns: a paleontological perspective. In: Ranker TA, Haufler CH (eds) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge, pp 332–366CrossRefGoogle Scholar
  138. Rouhan G, Dubuisson J-Y, Rakotondrainibe F, Motley TJ, Mickel JT, Labat J-N, Moran RC (2004) Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: contributions of species from the Indian Ocean area. Mol Phylogenet Evol 33:745–763PubMedCrossRefGoogle Scholar
  139. Rydin C, Wikström N, Wikstrom N (2002) Phylogeny of Isoëtes (Lycopsida): resolving basal relationships using rbcL sequences. Taxon 51:83CrossRefGoogle Scholar
  140. Sanchez-Baracaldo P (2004) Phylogenetic relationships of the subfamily Taenitidoideae, Pteridaceae. Am Fern J 94:126–142CrossRefGoogle Scholar
  141. Schneider H, Russell SJ, Cox CJ, Bakker F, Henderson S, Rumsey F, Barrett J, Gibby M, Vogel JC (2004a) Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL-F Spacer Sequences (Polypodiidae, Aspleniaceae) and its implications for biogeography. Syst Bot 29:260–274CrossRefGoogle Scholar
  142. Schneider H, Schuettpelz E, Pryer KM, Cranfill R, Magallón S, Lupia R (2004b) Ferns diversified in the shadow of angiosperms. Nature 428:553–557PubMedCrossRefGoogle Scholar
  143. Schneider H, Smith AR, Cranfill R, Hildebrand TJ, Haufler CH, Ranker TA (2004c) Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Mol Phylogenet Evol 31:1041–1063PubMedCrossRefGoogle Scholar
  144. Schneider H, Kreier HP, Hovenkamp P, Janssen T (2008) Phylogenetic relationships of the fern genus Christiopteris shed new light onto the classification and biogeography of drynarioid ferns. Bot J Linn Soc 157:645–656CrossRefGoogle Scholar
  145. Schneider H, Smith AR, Pryer KM (2009) Is morphology really at odds with molecules in estimating fern phylogeny? Syst Bot 34:455–475CrossRefGoogle Scholar
  146. Schuettpelz E, Hoot SB (2006) Inferring the root of Isoëtes: exploring alternatives in the absence of an acceptable outgroup. Syst Bot 31:258–270CrossRefGoogle Scholar
  147. Schuettpelz E, Pryer KM (2006) Reconciling extreme branch length differences: decoupling time and rate through the evolutionary history of filmy ferns. Syst Biol 55:485–502PubMedCrossRefGoogle Scholar
  148. Schuettpelz E, Pryer KM (2007) Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56:1037CrossRefGoogle Scholar
  149. Schuettpelz E, Pryer KM (2008) Fern phylogeny. In: Ranker TA, Haufler CH (eds) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge, pp 395–416CrossRefGoogle Scholar
  150. Schuettpelz E, Pryer KM (2009) Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy. Proc Natl Acad Sci U S A 106:11200–11205PubMedPubMedCentralCrossRefGoogle Scholar
  151. Schuettpelz E, Korall P, Pryer KM (2006) Plastid atpA data provide improved support for deep relationships among ferns. Taxon 55:897–906CrossRefGoogle Scholar
  152. Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM (2007) A molecular phylogeny of the fern family Pteridaceae: assessing overall relationships and the affinities of previously unsampled genera. Mol Phylogenet Evol 44:1172–1185PubMedCrossRefGoogle Scholar
  153. Schuettpelz E, Pryer KM, Windham MD (2015) A unified approach to taxonomic delimitation in the fern genus Pentagramma (Pteridaceae). Syst Bot 40:629–644CrossRefGoogle Scholar
  154. Schuettpelz E, Chen C-W, Kessler M, Pinson JB, Johnson G, Davila A, Cochran AT, Huiet L, Pryer KM (2016) A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data. Taxon 65:708–722CrossRefGoogle Scholar
  155. Sessa EB, Zimmer EA, Givnish TJ (2012) Phylogeny, divergence times, and historical biogeography of New World Dryopteris (Dryopteridaceae). Am J Bot 99:730–750PubMedCrossRefGoogle Scholar
  156. Sessa EB, Banks JA, Barker MS, Der JP, Duffy AM, Graham SW, Hasebe M, Langdale J, Li F-W, Marchant DB, Pryer KM, Rothfels CJ, Roux SJ, Salmi ML, Sigel EM, Soltis DE, Soltis PS, Stevenson DW, Wolf PG (2014) Between two fern genomes. GigaScience 3:15PubMedPubMedCentralCrossRefGoogle Scholar
  157. Shao Y, Wei R, Zhang X, Xiang Q (2015) Molecular phylogeny of the cliff ferns (Woodsiaceae: Polypodiales) with a proposed infrageneric classification. Plos One 10:e0136318PubMedPubMedCentralCrossRefGoogle Scholar
  158. Sigel EM, Windham MD, Huiet L, Yatskievych G, Pryer KM (2011) Species relationships and farina evolution in the cheilanthoid fern genus Argyrochosma (Pteridaceae). Syst Bot 36:554–564CrossRefGoogle Scholar
  159. Skog JE (2001) Biogeography of Mesozoic leptosporangiate ferns related to extant ferns. Brittonia 53:236–269CrossRefGoogle Scholar
  160. Skog JE, Banks HP (1973) Ibyka amphikoma, gen. et sp. n., a new protoarticulate precursor from the late Middle Devonian of New York State. Am J Bot 60:366–380CrossRefGoogle Scholar
  161. Skog JE, Zimmer EA, Mickel JT (2002) Additional support for two subgenera of Anemia (Schizaeaceae) from data for the chloroplast intergenic spacer region trnL-F and morphology. Am Fern J 92:119CrossRefGoogle Scholar
  162. Smith AR, Cranfill RB (2002) Intrafamilial relationships of the Thelypteroid Ferns (Thelypteridaceae). Am Fern J 92:131CrossRefGoogle Scholar
  163. Smith AR, Tuomisto H, Pryer KM, Hunt JS, Wolf PG (2001) Metaxya lanosa, a second species in the genus and fern family Metaxyaceae. Syst Bot 26:480–486Google Scholar
  164. Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG (2006) A classification for extant ferns. Taxon 55:705–731CrossRefGoogle Scholar
  165. Stanich NA, Rothwell GW, Stockey RA (2009) Phylogenetic diversification of Equisetum (Equisetales) as inferred from lower cretaceous species of British Columbia, Canada. Am J Bot 96:1289–1299PubMedCrossRefGoogle Scholar
  166. Stockey RA, Rothwell GW (2006) Introduction: evolution of modern ferns. Int J Plant Sci 167:613–614CrossRefGoogle Scholar
  167. Sundue M (2012) Galactodenia, a new genus of grammitid ferns segregated from Terpsichore (Polypodiaceae). Subterranean Biol 37Google Scholar
  168. Sundue MA, Islam MB, Ranker TA (2010) Systematics of grammitid ferns (Polypodiaceae): using morphology and plastid sequence data to resolve the circumscriptions of Melpomene and the polyphyletic genera Lellingeria and Terpsichore. Syst Bot 35:701–715CrossRefGoogle Scholar
  169. Sundue MA, Parris BS, Ranker TA, Smith AR, Fujimoto EL, Zamora-Crosby D, Morden CW, Chiou W-L, Chen C-W, Rouhan G, Hirai RY, Prado J (2014) Global phylogeny and biogeography of grammitid ferns (Polypodiaceae). Mol Phylogenet Evol 81:195–206PubMedCrossRefGoogle Scholar
  170. Taylor WC, Hickey RJ (1992) Habitat, evolution, and speciation in Isoetes. Ann Mo Bot Gard 79:613CrossRefGoogle Scholar
  171. Testo W, Sundue M (2016) A 4000-species dataset provides new insight into the evolution of ferns. Mol Phylogenet Evol 105:200–211PubMedCrossRefGoogle Scholar
  172. Tidwell WD, Ash SR (1994) A review of selected triassic to early Cretaceous ferns. J Plant Res 107:417–442CrossRefGoogle Scholar
  173. Tomescu AMF (2009) Megaphylls, microphylls and the evolution of leaf development. Trends Plant Sci 14:5–12PubMedCrossRefGoogle Scholar
  174. Vangerow S, Teerkorn T, Knoop V (1999) Phylogenetic information in the mitochondrial nad5 gene of pteridophytes: RNA editing and intron sequences. Plant Biol 1:235–243CrossRefGoogle Scholar
  175. Vanneste K, Sterck L, Myburg AA, Van de Peer Y, Mizrachi E (2015) Horsetails are ancient polyploids: evidence from Equisetum giganteum. Plant Cell 27:1567–1578PubMedPubMedCentralCrossRefGoogle Scholar
  176. Vasco A, Moran RC, Rouhan G (2009) Circumscription and phylogeny of the Elaphoglossum ciliatum group (E. sect. Lepidoglossa, Dryopteridaceae) based on cpDNA sequences. Taxon 58:825–834Google Scholar
  177. Vasco A, Moran RC, Ambrose BA (2013) The evolution, morphology, and development of fern leaves. Front Plant Sci 4:345PubMedPubMedCentralCrossRefGoogle Scholar
  178. Vasco A, Lóriga J, Rouhan G, Ambrose BA, Moran RC (2015) Divided leaves in the genus Elaphoglossum (Dryopteridaceae): a phylogeny of Elaphoglossum section Squamipedia. Syst Bot 40:46–55CrossRefGoogle Scholar
  179. Vasco A, Smalls TL, Graham SW, Cooper ED, Wong GK-S, Stevenson DW, Moran RC, Ambrose BA (2016) Challenging the paradigms of leaf evolution: class III HD-Zips in ferns and lycophytes. New Phytol 212:745–758PubMedCrossRefGoogle Scholar
  180. Volin JC, Lott MS, Muss JD, Owen D (2004) Predicting rapid invasion of the florida everglades by old world climbing fern (Lygodium microphyllum). Divers Distrib 10:439–446CrossRefGoogle Scholar
  181. Wagner WH, Beitel JM (1992) Generic classification of modern North American Lycopodiaceae. Ann Mo Bot Gard 79:676CrossRefGoogle Scholar
  182. Watkins JE, Cardelús CL (2012) Ferns in an angiosperm world: cretaceous radiation into the epiphytic niche and diversification on the forest floor. Int J Plant Sci 173:695–710CrossRefGoogle Scholar
  183. Wei R, Schneider H, Zhang XC (2013) Toward a new circumscription of the twinsorus-fern genus Diplazium (Athyriaceae): a molecular phylogeny with morphological implications and infrageneric taxonomy. Taxon 62:441–457CrossRefGoogle Scholar
  184. Wei R, Xiang Q, Schneider H, Sundue MA, Kessler M, Kamau PW, Hidayat A, Zhang X (2015) Eurasian origin, boreotropical migration and transoceanic dispersal in the pantropical fern genus Diplazium (Athyriaceae). J Biogeogr 42:1809–1819CrossRefGoogle Scholar
  185. Weststrand S, Korall P (2016a) Phylogeny of Selaginellaceae: there is value in morphology after all! Am J Bot 103:2136–2159PubMedCrossRefGoogle Scholar
  186. Weststrand S, Korall P (2016b) A subgeneric classification of Selaginella (Selaginellaceae). Am J Bot 103:2160–2169PubMedCrossRefGoogle Scholar
  187. Whitten WM, Jacono CC, Nagalingum NS (2012) An expanded plastid phylogeny of Marsilea with emphasis on North American species. Am Fern J 102:114–135CrossRefGoogle Scholar
  188. Wickett NJ, Mirarab S, Nguyen N, Warnow T, Carpenter E, Matasci N, Ayyampalayam S, Barker MS, Burleigh JG, Gitzendanner MA, Ruhfel BR, Wafula E, Der JP, Graham SW, Mathews S, Melkonian M, Soltis DE, Soltis PS, Miles NW, Rothfels CJ, Leebens-Mack J (2014) Phylotranscriptomic analysis of the origin and early diversification of land plants. Proc Natl Acad Sci U S A 111:E4859–E4868PubMedPubMedCentralCrossRefGoogle Scholar
  189. Wikström N, Kenrick P (2001) Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by use of nonparametric rate smoothing. Mol Phylogenet Evol 19:177–186PubMedCrossRefGoogle Scholar
  190. Wikström N, Pryer KM (2005) Incongruence between primary sequence data and the distribution of a mitochondrial atp1 group II intron among ferns and horsetails. Mol Phylogenet Evol 36:484–493PubMedCrossRefGoogle Scholar
  191. Wikström N, Kenrick P, Vogel JC (2002) Schizaeaceae: a phylogenetic approach. Rev Palaeobot Palynol 119:35–50CrossRefGoogle Scholar
  192. Wolf PG (1997) Evaluation of atpB nucleotide sequences for phylogenetic studies of ferns and other pteridophytes. Am J Bot 84:1429–1440PubMedCrossRefGoogle Scholar
  193. Yatabe Y, Nishida H, Murakami N (1999) Phylogeny of osmundaceae inferred from rbcL nucleotide sequences and comparison to the fossil evidences. J Plant Res 112:397–404CrossRefGoogle Scholar
  194. Zhang LB, Zhang L (2012) The inclusion of Acrophorus, Diacalpe, Nothoperanema, and Peranema in Dryopteris: the molecular phylogeny, systematics, and nomenclature of Dryopteris subg. Nothoperanema (Dryopteridaceae). Taxon 61:1199–1216Google Scholar
  195. Zhang L-B, Zhang L (2015) Didymochlaenaceae: a new fern family of eupolypods I (Polypodiales). Taxon 64:27–38CrossRefGoogle Scholar
  196. Zhang L-B, Zhang L, Dong S-Y, Sessa EB, Gao X-F, Ebihara A (2012) Molecular circumscription and major evolutionary lineages of the fern genus Dryopteris (Dryopteridaceae). BMC Evol Biol 12:180PubMedPubMedCentralCrossRefGoogle Scholar
  197. Zhang L, Rothfels CJ, Ebihara A, Schuettpelz E, Le Péchon T, Kamau P, He H, Zhou X-M, Prado J, Field A, Yatskievych G, Gao X-F, Zhang L-B (2015) A global plastid phylogeny of the brake fern genus Pteris (Pteridaceae) and related genera in the Pteridoideae. Cladistics Int J Willi Hennig Soc 31:406–423CrossRefGoogle Scholar
  198. Zhang L, Schuettpelz E, Rothfels CJ, Zhou X-M, Gao X-F, Zhang L-B (2016) Circumscription and phylogeny of the fern family Tectariaceae based on plastid and nuclear markers, with the description of two new genera: Draconopteris and Malaifilix (Tectariaceae). Taxon 65:723–738CrossRefGoogle Scholar
  199. Zhou X-M, Zhang L-B (2015) A classification of Selaginella (Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features. Taxon 64:1117–1140CrossRefGoogle Scholar
  200. Zhou X, Rothfels CJ, Zhang L, He Z, Le Péchon T, He H, Lu NT, Knapp R, Lorence D, He X, Gao X, Zhang L (2016) A large-scale phylogeny of the lycophyte genus Selaginella (Selaginellaceae: Lycopodiopsida) based on plastid and nuclear loci. Cladistics Int J Willi Hennig Soc 32:360–389CrossRefGoogle Scholar

Copyright information

© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Department of BiologyUniversity of FloridaGainesvilleUSA

Personalised recommendations