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Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE

Cytological study of Polystichum (Dryopteridaceae) species from southern South America

Rita E. Morero A B D , Franco E. Chiarini A , Juan Urdampilleta A , Gloria E. Barboza A B and David S. Barrington C
+ Author Affiliations
- Author Affiliations

A Instituto Multidisciplinario de Biología Vegetal (IMBIV-CONICET), and Universidad Nacional de Córdoba CC 495, CP 5000, Córdoba, Argentina.

B Facultad de Ciencias Químicas, Universidad Nacional de Córdoba, Haya de la Torre y Medina Allende, Córdoba, Argentina.

C University of Vermont, Pringle Herbarium, Torrey Hall, 27 Colchester Avenue, Burlington, VT 05405, USA.

D Corresponding author. Email: ritamorero@gmail.com

Australian Journal of Botany 63(5) 403-414 https://doi.org/10.1071/BT14287
Submitted: 24 October 2014  Accepted: 25 February 2015   Published: 18 May 2015

Abstract

Polystichum is one of the most diverse genera of ferns, with 360–400 species distributed worldwide. South America harbors ~40 species, clustered in three centres of diversity, namely, the Northern and Central Andes Center (NCC), the Brazilian Center (BC) and the Southern South America Center (SSC). To increase our understanding of the systematic relationships within Polystichum, mitotic chromosomes and spore features were studied in nine species from Argentina and Chile. All species presented the basic number x = 41, with different ploidy levels (2x, 4x and 8x). In general, chromosomes were homogeneous in size (average length 2.50–5.75 µm) and mostly subtelocentric; centromeres were inconspicuous and secondary constrictions were frequently observed. All species presented 64 spores per sporangium, suggesting normal sexual reproduction. Significant differences in spore size were found among species and it was positively correlated with ploidy level. A relationship between sum total chromosome length and ploidy level was observed. However, there was also a reduction in single-chromosome length in the polyploids, pointing to genome downsizing. Our results agree with previous records, with diploids being frequent among NCC species and absent among SSC species. In addition to sharing very specific morphological characters, SSC species are cytologically characterised by being polyploids (4x and 8x). A literature survey covering 116 species of Polystichum revealed that Australian and New Zealand Polystichum species exhibit similarly high frequencies of polyploidy. In the case of P. tetragonum (2n = 164), endemic to the Juan Fernandez archipelago, our data suggested that it was originated by transoceanic migration from a South American ancestor, probably also tetraploid.

Additional keywords: chromosome number, polyploidy, spore size.


References

Abraham A, Ninan CA, Mathew PM (1962) Studies on the cytology and phylogeny of the pteridophytes. VII. Observations on one hundred species of south Indian ferns. The Journal of the Indian Botanical Society 41, 339–421.

Barker MS, Wolf PG (2010) Unfurling fern biology in the genomics age. Bioscience 60, 177–185.
Unfurling fern biology in the genomics age.Crossref | GoogleScholarGoogle Scholar |

Barrington DS (1985a) Hybridization in Costa Rican Polystichum. Proceedings of the Royal Society of Edinburgh 86B, 335–340.

Barrington DS (1985b) The present evolutionary and taxonomic status of the fern genus Polystichum: the 1984 Botanical Society of America Pteridophyte, section symposium. American Fern Journal 75, 22–28.
The present evolutionary and taxonomic status of the fern genus Polystichum: the 1984 Botanical Society of America Pteridophyte, section symposium.Crossref | GoogleScholarGoogle Scholar |

Barrington DS (1985c) The morfophology and origin of a new Polystichum hybrid from Costa Rica. Systematic Botany 10, 199–204.
The morfophology and origin of a new Polystichum hybrid from Costa Rica.Crossref | GoogleScholarGoogle Scholar |

Barrington DS (1990) Hibridization and allopolyploidy in Central American Polystichum: citological and isozyme documentation. Annals of the Missouri Botanical Garden 77, 297–305.
Hibridization and allopolyploidy in Central American Polystichum: citological and isozyme documentation.Crossref | GoogleScholarGoogle Scholar |

Barrington DS (2003) Polystichum lilianae sp. nov. (Dryopteridaceae) and its relationships to P. fournieri and P. turrialbae. Brittonia 55, 317–325.
Polystichum lilianae sp. nov. (Dryopteridaceae) and its relationships to P. fournieri and P. turrialbae.Crossref | GoogleScholarGoogle Scholar |

Barrington DS (2006) ‘Cytological information for the genus Polystichum.’ (University of Vermont, Burlington, VT) Available at http://www.uvm.edu/~dbarring/polycytology.pdf. [Verified 14 November 2014]

Barrington DS, Driscoll HE (2005) ‘Minisymposium on the fern genus Polystichum.’ (British Pteridological Society, Kew: London) Available at http://www.uvm.edu/~dbarring/kewposter.pdf. [Verified 14 December 2014]

Barrington DS, Paris CA, Ranker TA (1986) Systematic inferences from spore and stomate size in the ferns. American Fern Journal 76, 149–159.
Systematic inferences from spore and stomate size in the ferns.Crossref | GoogleScholarGoogle Scholar |

Barrington DS, Haufler CH, Werth CR (1989) Hybridization, reticulation and species concepts in the ferns. American Fern Journal 79, 55–64.
Hybridization, reticulation and species concepts in the ferns.Crossref | GoogleScholarGoogle Scholar |

Bennett MD, Leitch IJ (2001) Nuclear DNA amounts in pteridophytes. Annals of Botany 87, 335–345.
Nuclear DNA amounts in pteridophytes.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3MXht1artbc%3D&md5=7da1505022460d15bc30e52f0c01f539CAS |

Bennett MD, Leitch IJ (2011) Nuclear DNA amounts in angiosperms: targets, trends and tomorrow. Annals of Botany 107, 467–590.
Nuclear DNA amounts in angiosperms: targets, trends and tomorrow.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3MXis1egtbo%3D&md5=1231e9e42a8840d500f8ae8b8ed99609CAS | 21257716PubMed |

Carr GD (1998) Chromosome evolution and speciation in Hawaiian flowering plants. In ‘Evolution and speciation of island plants’. (Eds TF Stuessy, M Ono) pp. 5–47. (Cambridge University Press: Cambridge, UK)

Condack JPS (2012) Filogenia de Polystichum (Dryopteridaceae) e revisão taxonômica das espécies ocorrentes no Brasil. [Phylogeny of Polystichum (Dryopteridaceae) and taxonomic revision of the species occurring in Brazil] Unpublished Dissertation Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro.

Condack JPS, Barrington DS, Sylvestre L (2010) ‘Phylogeny and biogeography of Polystichum Roth. (Dryopteridaceae) with emphasis on the species occurring at the Brazilian center of diversity’. Botany 2010 conference, Providence, RI. Available at http://2010.botanyconference.org/engine/search/index.php?func=detailandaid=1120. [Verified 31 December 2014]

Condack JPS, McHenry MA, Morero RE, Sylvestre LS, Barrington DS (2013) Polystichum montevidense demystified: molecular and morphologic data reveal a cohesive, widespread South American species. American Fern Journal 103, 118–130.
Polystichum montevidense demystified: molecular and morphologic data reveal a cohesive, widespread South American species.Crossref | GoogleScholarGoogle Scholar |

Crane FW (1953) Spore studies in Dryopteris, I. American Fern Journal 43, 159–169.
Spore studies in Dryopteris, I.Crossref | GoogleScholarGoogle Scholar |

Crawford DJ, Stuessy TF, Cosner MB, Haines DW, Silva M, Baeza M (1992) Evolution of the genus Dendroseris (Asteraceae, Lactuceae) on the Juan Fernandez Islands: evidence from chloroplast and ribosomal DNA. Systematic Botany 17, 676–682.
Evolution of the genus Dendroseris (Asteraceae, Lactuceae) on the Juan Fernandez Islands: evidence from chloroplast and ribosomal DNA.Crossref | GoogleScholarGoogle Scholar |

Crawford DJ, Sang T, Stuessy TF, Kim S-C, Silva M (1998) Dendroseris (Asteraceae: Lactuceae) and Robinsonia (Asteraceae: Senecioneae) on the Juan Fernández Islands: similarities and differences in biology and phylogeny. In ‘Evolution and speciation of island plants’. (Eds TF Stuessy, M Ono) pp. 97–119 (Cambridge University Press: Cambridge, UK)

Cubas P, Pardo C (1992) Perispore structure in Polystichum setiferum, P. aculeatum and their hybrid P. xbicknellii. American Fern Journal 82, 125–128.
Perispore structure in Polystichum setiferum, P. aculeatum and their hybrid P. xbicknellii.Crossref | GoogleScholarGoogle Scholar |

Daigobo S (1972) Taxonomical studies on the fern genus Polystichum in Japan, Ryukyu, and Taiwan. Science Reports of the Tokyo Kyoiku Daigaku. Section B 15, 57–80.

Dart S, Kron P, Mable BK (2004) Characterizing polyploidy in Arabidopsis lyrata using chromosome counts and flow cytometry. Canadian Journal of Botany 82, 185–197.
Characterizing polyploidy in Arabidopsis lyrata using chromosome counts and flow cytometry.Crossref | GoogleScholarGoogle Scholar |

Dawson MI, Brownsey PJ, Lovis JD (2000) Index of chromosome numbers of indigenous New Zealand pteridophytes. New Zealand Journal of Botany 38, 25–46.
Index of chromosome numbers of indigenous New Zealand pteridophytes.Crossref | GoogleScholarGoogle Scholar |

de la Sota ER, Giúdice GE, Ponce M, Ramos Giacosa JP, Arturi M (2004) Relaciones fitogeográficas de la flora pteridofítica serrana bonaerense. Boletín de la Sociedad Argentina de Botánica 39, 181–194.

Devi S (1977) Scanning electron microscope observations of the spores of some species of Polystichum. Review of Palaeobotany and Palynology 23, 139–144.
Scanning electron microscope observations of the spores of some species of Polystichum.Crossref | GoogleScholarGoogle Scholar |

Di Rienzo JA, Balzarini MG, González L, Tablada M, Guzmán W, Robledo CW, Casanoves F (2002) ‘Software InfoStat versión 1.1.’ (Grupo InfoStat, F. Cs. Agrarias, Universidad Nacional de Córdoba: Argentina) Available at http://www.infostat.com.ar. [Verified 30 November 2014]

Dimitrova D, Greilhuber J (2000) Karyotype and DNA-content evolution in ten species of Crepis (Asteraceae) distributed in Bulgaria. Botanical Journal of the Linnean Society 132, 281–297.
Karyotype and DNA-content evolution in ten species of Crepis (Asteraceae) distributed in Bulgaria.Crossref | GoogleScholarGoogle Scholar |

Fraser-Jenkins CR (1997) ‘Himalayan ferns: a guide to Polystichum.’ (International Booksellers: Dehradun, India)

Garnatje T, Valles J, Garcia S, Hidalgo O, Sanz M, Canela MA, Siljak-Yakovlev S (2004) Genome size in Echinops L. and related genera (Asteraceae, Cardueae): karyological, ecological and phylogenetic implications. Biology of the Cell 96, 117–124.
Genome size in Echinops L. and related genera (Asteraceae, Cardueae): karyological, ecological and phylogenetic implications.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXisF2isbY%3D&md5=7f131ebf1dd19b92899c8fcb166253c2CAS | 15050366PubMed |

Gastony GJ (1991) Gene silencing in a polyploid homosporous fern: paleopolyploidy revisited. Proceedings of the National Academy of Sciences, USA 88, 1602–1605.
Gene silencing in a polyploid homosporous fern: paleopolyploidy revisited.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3MXhsFygt7Y%3D&md5=947b6c4962433166f236d8b964f16d2eCAS |

Guillén RH, Daviña JR (2005) Chromosome studies in species of Asplenium (Aspleniaceae) from Argentina. Darwiniana 43, 44–51.

Haufler CH, Soltis DE (1986) Genetic evidence indicates that homosporous ferns with high chromosome numbers may be diploid. Proceedings of the National Academy of Sciences, USA 83, 4389–4393.
Genetic evidence indicates that homosporous ferns with high chromosome numbers may be diploid.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL28Xktl2msbs%3D&md5=33e41167d69f8a8a1846d32f05b1addfCAS |

Hijmans RJ, Guarino L, Bussink C, Mathur P, Cruz M, Barrentes I, Rojas E (2004) ‘DIVA-GIS. V 5. A geographic information system for the analysis of species distribution data.’ Available at http://www.diva-gis.org

Ivanova D, Piekos-Mirkowa H (2003) Chromosome numbers of Polish ferns. Acta Biologica Cracoviensia. Series; Botanica 45, 93–99.

Jara-Seguel P, Romero-Mieres M, Palma-Rojas C (2006) Chromosome numbers of Chilean pteridophytes: first contribution. Gayana Botánica 63, 115–118.

Kato M, Nakato N (1999) A cytotaxonomic study of Hainan (S China) pteridophytes with notes on polyploidy and apogamy of Chinese species. In ‘Ching Memorial Volume’. (Eds XC Zhang, KH Shing) pp. 1–19. (Chinese Forestry Publishing House: Beijing)

Khullar SP, Gupta SC (1978) Cytotaxonomy of the genus Polystichum in the western Himalayas. Plant Systematics and Evolution 129, 269–275.
Cytotaxonomy of the genus Polystichum in the western Himalayas.Crossref | GoogleScholarGoogle Scholar |

Khullar SP, Sharma SS, Verma SC (1988) SOCGI plant chromosome number reports. VI. Journal of Cytology and Genetics 23, 38–52.

Klekowski EJ (1973) Sexual and subsexual systems in homosporous pteridophytes: a new hypothesis. American Journal of Botany 60, 535–544.
Sexual and subsexual systems in homosporous pteridophytes: a new hypothesis.Crossref | GoogleScholarGoogle Scholar |

Klekowski EJ, Baker HG (1966) Evolutionary significance of polyploidy in the Pteridophyta. Science 153, 305–307.
Evolutionary significance of polyploidy in the Pteridophyta.Crossref | GoogleScholarGoogle Scholar | 17780002PubMed |

Knobloch I (1996) ‘Pteridophyte hybrids and their derivatives.’ (Michigan State University Department of Botany and Plant Pathology: East Lansing, MI)

Leitch IJ, Bennett MD (2004) Genome downsizing in polyploid plants. Biological Journal of the Linnean Society. Linnean Society of London 82, 651–663.
Genome downsizing in polyploid plants.Crossref | GoogleScholarGoogle Scholar |

Leitch IJ, Soltis DE, Soltis PS, Bennett MD (2005) Evolution of DNA amounts across land plants (Embryophyta). Annals of Botany 95, 207–217.
Evolution of DNA amounts across land plants (Embryophyta).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXht1KiurY%3D&md5=5f435e58e737f539ebda786047bb57edCAS | 15596468PubMed |

Levin DA (2002) ‘The role of chromosomal change in plant evolution.’ (Oxford University Press: New York)

Li CS, Lu SG, Yang Q (2004) Asian origin for Polystichum (Dryopteridaceae) based on rbcL sequences. Chinese Science Bulletin 49, 1146–1150.
Asian origin for Polystichum (Dryopteridaceae) based on rbcL sequences.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXmvVCqtbs%3D&md5=c9352110b7cb107b1dbedf7109e1cccaCAS |

Li C-X, Lu SG, Barrington DS (2008) Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL–Fanbrps4–trnS). Journal of Plant Research 121, 19–26.
Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL–Fanbrps4–trnS).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXktVOnuw%3D%3D&md5=b4d5cdcc7e3825199bdf8937edfa2192CAS | 18000642PubMed |

Lin S-J, Ebihara A, Wang D, Iwatsuki K (2011) Systematic studies of Polystichum (Dryopteridaceae) in Japan (I): P. fibrillosopaleaceum var. marginale is a diploid hybrid between P. fibrilloso paleaceum and P. igaense. American Fern Journal 101, 295–306.
Systematic studies of Polystichum (Dryopteridaceae) in Japan (I): P. fibrillosopaleaceum var. marginale is a diploid hybrid between P. fibrilloso paleaceum and P. igaense.Crossref | GoogleScholarGoogle Scholar |

Little DP, Barrington DS (2003) Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). American Journal of Botany 90, 508–514.
Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae).Crossref | GoogleScholarGoogle Scholar | 21659143PubMed |

Löve A, Löve D, Pichi Sermolli REG (1977) ‘Cytotaxonomical Atlas of the Pteridophyta.’ (J Cramer: Vaduz)

Lu J-M, Li D-Z, Wu D (2006a) Chromosome numbers of four genera in the Dryopteridaceae. Acta Phytotaxonomica Sinica 44, 516–522.
Chromosome numbers of four genera in the Dryopteridaceae.Crossref | GoogleScholarGoogle Scholar |

Lu J-M, Cheng X, Wuand D, Li D-Z (2006b) Chromosome study of the fern genus Cyrtomium (Dryopteridaceae). Botanical Journal of the Linnean Society 150, 221–228.
Chromosome study of the fern genus Cyrtomium (Dryopteridaceae).Crossref | GoogleScholarGoogle Scholar |

Manton I (1950) ‘Problems of cytology and evolution in the Pteridophyta.’ (Cambridge University Press: Cambridge, UK)

Marcon AB, Barros ICL, Guerra M (2003a) A karyotype comparison between two closely related Acrostichum L. (Pteridaceae) species. American Fern Journal 93, 116–125.
A karyotype comparison between two closely related Acrostichum L. (Pteridaceae) species.Crossref | GoogleScholarGoogle Scholar |

Marcon AB, Barros ICL, Guerra M (2003b) Cariologia de algumas espécies de pteridófitas ocorrentes no nordeste do Brasil. Acta botânica Brasilica 17, 19–26.
Cariologia de algumas espécies de pteridófitas ocorrentes no nordeste do Brasil.Crossref | GoogleScholarGoogle Scholar |

Marcon AB, Barros ICL, Guerra M (2005) Variation in chromosome numbers, CMA Bands and 45S rDNA sites in species of Selaginella (Pteridophyta). Annals of Botany 95, 271–276.
Variation in chromosome numbers, CMA Bands and 45S rDNA sites in species of Selaginella (Pteridophyta).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXht1KitLc%3D&md5=204a8d71ad1967e7ef015ac40aa4bc43CAS | 15567808PubMed |

McHenry MA, Barrington DS (2014) Phylogeny and biogeography of exindusiate Andean Polystichum (Dryopteridaceae). American Journal of Botany 101, 365–375.
Phylogeny and biogeography of exindusiate Andean Polystichum (Dryopteridaceae).Crossref | GoogleScholarGoogle Scholar | 24525330PubMed |

Medina FJ, Solanilla EL, Sánchez-Pina MA, Fernández-Gómez ME, Risueño MC (1986) Cytological approach to the nucleolar functions detected by silver-staining. Chromosoma 94, 259–266.
Cytological approach to the nucleolar functions detected by silver-staining.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL2sXhsF2huw%3D%3D&md5=11523a4e310cf8661a344e6a3f24eae9CAS |

Mitui K (1973) Characteristics of the perispore of the genus Polystichum by the scanning electron microscope. Bulletin of Nippon Dental College. General Education 2, 105–123.

Miyamoto F, Nakamura T (1983) Tree new hybrids of Polystichum from Taiwan. Japanese Journal of Botany 58, 146–150.

Moran RC (1982) The Asplenium trichomanes complex in the United States and adjacent Canada. American Fern Journal 72, 5–11.
The Asplenium trichomanes complex in the United States and adjacent Canada.Crossref | GoogleScholarGoogle Scholar |

Morbelli MA (1980) Morfología de las esporas de Pteridophyta presentes en la región Fuego-Patagónica República Argentina. Opera lilloana 28, 1–138.

Morero R, Barrington DS, Barboza G, Chiapella JO (2013) Filogenia preliminar del grupo Sudamericano Austral de Polystichum (Dryopteridaceae) de Chile y Argentina. Boletín de la Sociedad Argentina de Botánica 48, 267

Nagl W, Ehrendorfer F (1974) DNA content, heterochromatin, mitotic index and growth in perennial and annual Anthemideae (Asteraceae). Plant Systematics and Evolution 123, 35–54.
DNA content, heterochromatin, mitotic index and growth in perennial and annual Anthemideae (Asteraceae).Crossref | GoogleScholarGoogle Scholar |

Otto SP (2007) The evolutionary consequences of polyploidy. Cell 131, 452–462.
The evolutionary consequences of polyploidy.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXhtlWitr%2FJ&md5=8fffc06d4b085c7faff8a285971af969CAS | 17981114PubMed |

Ozkan H, Tuna M, Arumuganathan K (2003) Non additive changes in genome size during allopolyploidization in the wheat (Aegilops–Triticum) group. The Journal of Heredity 94, 260–264.
Non additive changes in genome size during allopolyploidization in the wheat (Aegilops–Triticum) group.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXkvVSmt70%3D&md5=a953bbaeb557524fe36e3dfab425f7ebCAS | 12816968PubMed |

Perrie LR, Brownsey PJ, Lockhart PJ, Brown EA, Large MF (2003a) Biogeography of temperate Australasian Polystichum ferns as inferred from chloroplast sequence and AFLP. Journal of Biogeography 30, 1729–1736.
Biogeography of temperate Australasian Polystichum ferns as inferred from chloroplast sequence and AFLP.Crossref | GoogleScholarGoogle Scholar |

Perrie LR, Brownsey PJ, Lockhart PJ, Large MF (2003b) Evidence for an allopolyploid complex in New Zealand Polystichum (Dryopteridaceae). New Zealand Journal of Botany 41, 189–215.
Evidence for an allopolyploid complex in New Zealand Polystichum (Dryopteridaceae).Crossref | GoogleScholarGoogle Scholar |

Ponce MM, Mehltreter K, de la Sota ER (2002) Análisis biogeográfico de la diversidad pteridofítica en Argentina y Chile continental. Revista Chilena de Historia Natural (Valparaiso, Chile) 75, 703–717.
Análisis biogeográfico de la diversidad pteridofítica en Argentina y Chile continental.Crossref | GoogleScholarGoogle Scholar |

Quintanilla LG, Escudero A (2006) Spore fitness components do not differ between diploid and allotetraploid species of Dryopteris (Dryopteridaceae). Annals of Botany 98, 609–618.
Spore fitness components do not differ between diploid and allotetraploid species of Dryopteris (Dryopteridaceae).Crossref | GoogleScholarGoogle Scholar | 16845140PubMed |

Rasband WS (2014) ‘ImageJ.’ (US National Institutes of Health: Bethesda, MD) Available at http://imagej.nih.gov/ij/. [Verified 31 December 2014]

Rodriguez R (1989) Phytogeographical and taxonomical comments on Chilean pteridophyta. Gayana Botánica 46, 199–208.

Roux JP (2000) The genus Polystichum (Dryopteridaceae) in Africa. Bulletin of the British Museum (Natural History). Botany 30, 33–79.

Roux JP (2001) A review of the fern genus Polystichum (Pteropsida: Dryopteridaceae) in Madagascar and the Mascarene Region. Adansonia 23, 265–287.

Roux JP (2004) The fern genus Polystichum (Pteropsida: Dryopteridaceae) in Macaronesia. Systematics and Biodiversity 2, 147–161.
The fern genus Polystichum (Pteropsida: Dryopteridaceae) in Macaronesia.Crossref | GoogleScholarGoogle Scholar |

Sanders RW, Stuessy TF, Rodriguez R (1983) Chromosome numbers from the flora of the Juan Fernandez Islands. American Journal of Botany 70, 799–810.
Chromosome numbers from the flora of the Juan Fernandez Islands.Crossref | GoogleScholarGoogle Scholar |

Shimura Y, Ooishi M (1980) A study on the chromosome numbers of some taxa of Japanese Polystichum. Taxon 28, 24–27.

Smith AR, Foster MS (1984) Chromosome numbers and ecological observations of ferns from El Tirol, Paraguay. The Fern Gazette 12, 321–329.

Smith AR, Mickel JT (1977) Chromosome counts for mexican ferns. Brittonia 29, 391–398.
Chromosome counts for mexican ferns.Crossref | GoogleScholarGoogle Scholar |

Soltis DE, Soltis PS (1999) Polyploidy: recurrent formation and genome evolution. Trends in Ecology & Evolution 14, 348–352.
Polyploidy: recurrent formation and genome evolution.Crossref | GoogleScholarGoogle Scholar |

Stergianou KK, Fowler K (1990) Chromosome numbers and taxonomic implications in the fern genus Azolla (Azollaceae). Plant Systematics and Evolution 173, 223–239.
Chromosome numbers and taxonomic implications in the fern genus Azolla (Azollaceae).Crossref | GoogleScholarGoogle Scholar |

Stuessy TF, Crawford DJ (1998). Chromosomal stasis during speciation in angiosperms of oceanic islands. In ‘Evolution and speciation of island plants’. (Eds TF Stuessy, M Ono) pp. 307–324. (Cambridge University Press: Cambridge, UK)

Stuessy TF, Sanders RW, Silva M (1984) Phytogeography and evolution of the flora of the Juan Fernandez Islands: a progress report. In ‘Biogeography of the tropical Pacific’. (Eds FJ Radovsky, PH Raven, SH Sohmer) pp. 55–69. (Association of Systematic Collections and BP Bishop Museum: Lawrence, KS)

Stuessy TF, Crawford DJ, Marticorena C (1990) Patterns of phylogeny in the endemic vascular flora of the Juan Fernandez Islands, Chile. Systematic Botany 15, 338–346.
Patterns of phylogeny in the endemic vascular flora of the Juan Fernandez Islands, Chile.Crossref | GoogleScholarGoogle Scholar |

Stuessy TF, Jakubowsky G, Salguero Gómez R, Pfosser M, Schlüter PM, Fer T, Sun B-Y, Kato H (2006) Anagenetic evolution in island plants. Journal of Biogeography 33, 1259–1265.
Anagenetic evolution in island plants.Crossref | GoogleScholarGoogle Scholar |

Taylor TMC, Lang F (1963) Chromosome counts in some British Columbia ferns. American Fern Journal 53, 123–126.
Chromosome counts in some British Columbia ferns.Crossref | GoogleScholarGoogle Scholar |

Tindale MD, Roy SK (2002) A cytotaxonomic survey of the Pteridophyta of Australia. Australian Systematic Botany 15, 839–937.
A cytotaxonomic survey of the Pteridophyta of Australia.Crossref | GoogleScholarGoogle Scholar |

Tryon AF, Lugardon B (1990) ‘Spore of the Pteridophyta: surface, wall structures and diversity based on electron microscope studies.’ (Springer-Verlag: New York)

Tsai JL, Shieh WC (1985) A cytotaxonomic survey of the fern family Aspidiaceae in Taiwan. Journal of Engineering Science NCHU 40, 121–144.

Von Kalm L, Smyth DR (1984) Ribosomal RNA genes and the substructure of nucleolar organizing regions in Lillium. Genome 26, 158–166.

Wagner WH (1962) Irregular morphological development in hybrid ferns. Phytomorphology 12, 87–100.

Wagner WH (1974) Structure of spores in relation to fern phylogeny. Annals of the Missouri Botanical Garden 61, 332–353.
Structure of spores in relation to fern phylogeny.Crossref | GoogleScholarGoogle Scholar |

Wagner DH (1979) Systematics of Polystichum in western North America and north of Mexico. Pteridología 1, 1–64.

Wagner FS (1980) New basic chromosome numbers for genera of neotropical ferns. American Journal of Botany 67, 733–738.
New basic chromosome numbers for genera of neotropical ferns.Crossref | GoogleScholarGoogle Scholar |

Wagner WH, Wagner FS, Taylor WC (1986) Detecting abortive spores in herbarium specimens of sterile hybrids. American Fern Journal 76, 129–140.
Detecting abortive spores in herbarium specimens of sterile hybrids.Crossref | GoogleScholarGoogle Scholar |

Warburton D, Henderson AS (1979) Sequential silver staining and hybridization in situ of nucleolus organizer regions in human cells. Cytogenetics and Cell Genetics 24, 168–175.
Sequential silver staining and hybridization in situ of nucleolus organizer regions in human cells.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaL3c%2FgtlSmtQ%3D%3D&md5=5dfacc50701afdc239bf692e6bc10933CAS |

Wendel JF (2000) Genome evolution in polyploids. Plant Molecular Biology 42, 225–249.
Genome evolution in polyploids.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXht1Grsrk%3D&md5=6f4920b37df8303cf5eca46d1d8063e3CAS | 10688139PubMed |

Werth CR, Windham MD (1991) A model for divergent, allopatric speciation of polyploidy pteridophytes resulting from silencing of duplicate-gene expression. American Naturalist 137, 515–526.
A model for divergent, allopatric speciation of polyploidy pteridophytes resulting from silencing of duplicate-gene expression.Crossref | GoogleScholarGoogle Scholar |

Xiang J-Y, Chengand X, Wu SG (2006) Chromosome numbers of 13 species in the genus Dryopteris (Dryopteridaceae) from Yunnan, China. Acta Phytotaxonomica Sinica 44, 304–319.
Chromosome numbers of 13 species in the genus Dryopteris (Dryopteridaceae) from Yunnan, China.Crossref | GoogleScholarGoogle Scholar |