The highest frequency of polyploidy among plants is considered to occur in the Pteridophytes. In this study, we focused on polyploidy displayed by a specific fern taxon, the genus Pteris L. (Pteridaceae), comprising over 250 species. Cytological data from 106 Pteris species were reviewed. The base number of chromosomes in Pteris is 29. Polyploids are frequently found in Pteris, including triploids, tetraploids, pentaploids, hexaploids, and octoploids. In addition, an aneuploid species, P. deltodon Bak., has been recorded. Furthermore, the relationship between polyploidy and reproductive biology is reviewed. Among these 106 Pteris species, 60% exhibit polyploidy: 22% show intraspecific polyploidy and 38% result from polyploid speciation. Apogamous species are common in Pteris. Diploids are the most frequent among Pteris species, and they can be sexual or apogamous. Triploids are apogamous; tetraploids are sexual or apogamous. Most Pteris species have one to two ploidy levels. The diverse ploidy levels suggest that these species have a complex evolutionary history and their taxonomic problems require further clarification. 1. Introduction Polyploidy provides a rapid route for species evolution and adaptation [1, 2]. Taxa arising from polyploidy are usually characterized by divers gene expression [3]. This variation in gene expression also has effects on ecological traits, which play an important role in speciation because a specialised niche is a key factor in the formation of new taxa [4–8]. For example, ecological isolation can allow taxa with genetic variation to become segregated [9]. It is estimated that the highest frequency of polyploidy is exhibited in ferns. The frequency of polyploid speciation in ferns is 31%, which is much higher than 15% in angiosperms [10]. In ferns, a special form of asexual reproduction known as apogamy is common [11, 12]. Apogamy provides a bypass to crossover mispairing of chromosomes and stabilises the reproduction of polyploids [13–15]. During metaphase I of meiosis, these polyploids present multivalents, which may have difficulty separating equally. Apogamous species are clonal hybrid genotypes, and, as a result, apogamy creates reproductive barriers that prevent gene flow among closely related taxa, thereby facilitating sympatric speciation [16]. Each taxon maintains an independent genetic lineage, leading eventually to a new species. Pteris L. (Pteridaceae) is a cosmopolitan fern genus with over 250 species. Some Pteris species have several different ploidy levels and are found in several geographical areas, such as
References
[1]
V. Grant, Plant Speciation, Columbia University Press, New York, NY, USA, 1981.
[2]
L. H. Rieseberg and J. H. Willis, “Plant speciation,” Science, vol. 317, no. 5840, pp. 910–914, 2007.
[3]
P. S. Soltis and D. E. Soltis, “The role of genetic and genomic attributes in the success of polyploids,” Proceedings of the National Academy of Sciences of the United States of America, vol. 97, no. 13, pp. 7051–7057, 2000.
[4]
C. N. Page, “Ecological strategies in fern evolution: a neopteridological overview,” Review of Palaeobotany and Palynology, vol. 119, no. 1-2, pp. 1–33, 2002.
[5]
C. H. Haufler, M. D. Windham, D. M. Britton, and S. J. Robinson, “Triploidy and its evolutionary significance in Cystopteris protrusa,” Canadian Journal of Botany, vol. 63, no. 10, pp. 1855–1863, 1985.
[6]
J. Ramsey and D. W. Schemske, “Neopolyploidy in flowering plants,” Annual Review of Ecology and Systematics, vol. 33, pp. 589–639, 2002.
[7]
D. M. Rosenthal, A. E. Schwarzbach, L. A. Donovan, O. Raymond, and L. H. Rieseberg, “Phenotypic differentiation between three ancient hybrid taxa and their parental species,” International Journal of Plant Sciences, vol. 163, no. 3, pp. 387–398, 2002.
[8]
B. L. Gross and L. H. Rieseberg, “The ecological genetics of homoploid hybrid speciation,” Journal of Heredity, vol. 96, no. 3, pp. 241–252, 2005.
[9]
J. C. Vogel, F. J. Rumsey, S. J. Russell et al., “Genetic structure, reproductive biology and ecology of isolated populations of Asplenium csikii (Aspleniaceae, Pteridophyta),” Heredity, vol. 83, no. 5, pp. 604–612, 1999.
[10]
T. E. Wood, N. Takebayashi, M. S. Barker, I. Mayrose, P. B. Greenspoon, and L. H. Rieseberg, “The frequency of polyploid speciation in vascular plants,” Proceedings of the National Academy of Sciences of the United States of America, vol. 106, no. 33, pp. 13875–13879, 2009.
[11]
I. Manton, Problems of Cytology and Evolution in the Pteridophyta, Columbia University Press, New York, NY, USA, 1950.
[12]
T. G. Walker, “Cytology and evolution in the fern genus Pteris L.,” Evolution, vol. 16, no. 1, pp. 17–43, 1962.
[13]
J. D. Lovis, “Evolutionary patterns and processes in ferns,” Advances in Botanical Research, vol. 4, no. C, pp. 229–415, 1978.
[14]
T. G. Walker, The Experimental Biology of Ferns, Academic Press, Londo, UK, 1979, Edited by A. F. Dyer.
[15]
L. G. Hickok and E. J. Klekowski Jr., “Inchoate speciation in Ceratopteris:an analysis od the synthesized hybrid C. richardii x C. pteridoides,” Evolution, vol. 28, pp. 439–446, 1974.
[16]
C. R. Werth and M. D. Windham, “A model for divergent, allopatric speciation of polyploid pteridophytes resulting from silencing of duplicate-gene expression,” The American Naturalist, vol. 137, no. 4, pp. 515–526, 1991.
[17]
P. Khare and S. Kaur, “Intraspecific polyploidy in Pteris vittata Linn.,” Cytologia, vol. 48, no. 1, pp. 21–25, 1983 (Japanese).
[18]
Z. R. Wang, “A preliminary study on cytology of Chinese Pteris,” Acta Phytotaxonomica Sinica, vol. 27, no. 6, pp. 421–438, 1989.
[19]
Y.-M. Huang, H.-M. Chou, J.-C. Wang, and W.-L. Chiou, “The distribution and habitats of the Pteris fauriei complex in Taiwan,” Taiwania, vol. 52, no. 1, pp. 49–58, 2007.
[20]
T. G. Walker, “The Pteris quadriaurita complex in Ceylon,” Kew Bulletin, vol. 14, no. 3, pp. 321–332, 1954.
[21]
T. G. Walker, “Hybridization in some species of Pteris L.,” Evolution, vol. 12, no. 1, pp. 82–92, 1958.
[22]
I. Manton, “Chromosomes and fern phylogeny with special reference to“Pteridaceae”,” Journal of the Linnean Society of London, vol. 56, no. 365, pp. 73–92, 1958.
[23]
K. Iwatsuki, lora of Japan-Pteridophyta and Gymnospermae, Kodansha, Tokyo, Japan, 1995, Edited by K. Iwatsuki, T. Yamazaki, D. E. Boufford and H. Ohba.
[24]
Index to Plant Chromosome Numbers (IPCN), Missouri Botanical Garden, St. Louis, Mo, USA, 1979, Edited by P. Goldblatt and D. E. Johnson, http://www.tropicos.org/Project/IPCN/.
[25]
T. G. Walker, “Additional cytogenetic notes on the pteridophytes of Jamaica,” Transactions of the Royal Society of Edinburgh, vol. 69, pp. 109–135, 1973.
[26]
L. S. Ammal and K. V. Bhavanandan, “Cytological studies on some members of Pteridaceae (sensu Copeland) from south India,” Indian Fern Journal, vol. 8, no. 1-2, pp. 87–92, 1991.
[27]
J.-L. Tsai and W.-C. Shieh, “A cytotaxonomic survey of the pteridophytes in Taiwan (2) chromosome and spore characteristics,” Journal of Science & Engineering, vol. 21, pp. 57–70, 1984.
[28]
Y. J. Chang, W. C. Shieh, and J. L. Tsai, “Studies on the karyotypes of the fern genus Pteris in Taiwan,” in Proceedings of the 2nd Seminar on Asian Pteridology, Taipei, Taiwan, 1992.
[29]
I. Manton and W. A. Sledge, “Observations on the cytology and taxonomy of the Pteridophyte flora of Ceylon,” Philosophical Transactions of the Royal Society B, vol. 238, pp. 127–185, 1954.
[30]
S. K. Roy and J. B. Singh, “A note on the chromosome numbers in some ferns from Pachmarhi Hills,” Central India, vol. 41, pp. 181–183, 1975.
[31]
P. N. Mehra, “Chromosome number in Himalayan ferns,” Research Bulletin (n.s.) of Panjab University, vol. 12, pp. 139–169, 1961.
[32]
Y. S. Chao, H. Y. Liu, Y. M. Huang, and W. L. Chiou, “Reproductive traits of Pteris cadieri and P. grevilleana in Taiwan: implications for their hybrid origin,” Botanical Studies, vol. 51, no. 2, pp. 209–216, 2010.
[33]
á. L?ve, D. L?ve, and R. E. G. Sermolli, Cytotaxonomical Atlas of the Pteridophyta, Vaduz, Liechtenstein, 1977, Edited by J. Cramer.
[34]
N. Nakat?, “A cytological study on an intermediate form between Pteris multifida and P. cretica,” Journal of Japanese Botany, vol. 50, no. 4, pp. 110–125, 1975.
[35]
R. P. Roy, B. M. B. Sinha, and A. R. Sakya, “Cytology of some ferns of Kathmandu valley,” Fern Gazette, vol. 10, pp. 193–199, 1971.
[36]
Y.-M. Huang, S.-Y. Hsu, T.-H. Hsieh, H.-M. Chou, and W.-L. Chiou, “Three Pteris species (Pteridaceae: Pteridophyta) reproduce by apogamy,” Botanical Studies, vol. 52, p. 79, 2011.
[37]
S. C. Verma and S. P. Khullar, “Cytogenetics of the Western Himalayan Pteris cretica complex,” Annals of Botany, vol. 29, no. 4, pp. 673–681, 1965.
[38]
S.-J. Lin, K. Iwatsuki, and M. Kato, “Cytotaxonomic study of ferns from China I. Species of Yunnan,” Journal of Japanese Botany, vol. 71, no. 4, pp. 214–222, 1996.
[39]
M. Kato, N. Nakato, X. Cheng, and K. Iwatsuki, “Cytotaxonomic study of ferns of Yunnan, southwestern China,” The Botanical Magazine Tokyo, vol. 105, no. 1, pp. 105–124, 1992.
[40]
W. H. J. Wagner and C. E. Nauman, “Pteris Xdelchampsii, a spontaneous fern hybrid from southern Florida,” American Fern Journal, vol. 72, pp. 97–102, 1982.
[41]
A. Abraham, “Studies on the cytology and phylogeny of the Pteridophytes VII. Observation on one hundred species of South Indian ferns,” Journal of the Indian Botanical Society, vol. 41, pp. 344–348, 1962.
[42]
P. I. Kuriachan and C. A. Ninan, Aspects of Plant Sciences, Today & Tomorrow's Printers, New Delhi, India, 1976, Edited by P. K. K. Nair.
[43]
W.-L. Chiou, “The gametophytes of Pteris ensiformis Burm,” in Proceedings of the Seminar of Asian Pteridology II, Taipei, Taiwan, 1992.
[44]
K. Mitui, “Chromosome numbers of some ferns in the Ryukyu Islands,” Journal of Japanese Botany, vol. 51, pp. 33–41, 1976.
[45]
N. Nakat?, “A cytogeographic study on the Japanese Pteris excelsa complex,” Journal of Japanese Botany, vol. 51, pp. 59–64, 1976.
[46]
Y. M. Huang, H. M. Chou, T. H. Hsieh, J. C. Wang, and W. L. Chiou, “Cryptic characteristics distinguish diploid and triploid varieties of Pteris fauriei (Pteridaceae),” Canadian Journal of Botany, vol. 84, no. 2, pp. 261–268, 2006.
[47]
N. Nakato, “Notes on chromosomes of Japanese pteridophytes,” Journal of Japanese Botany, vol. 65, pp. 204–209, 1990.
[48]
T. Suzuki and K. Iwatsuki, “Genetic variation in apogamous fern Pteris cretica L. in Japan,” Heredity, vol. 65, pp. 221–227, 1990.
[49]
N. Nakato, “Notes on chromosomes of Japanese pteridophytes,” Journal of Japanese Botany, vol. 63, pp. 214–218, 1988.
[50]
K. Mitui, “Chromosomes and speciation in ferns,” Science Reports of the Tokyo Kyoiku Daigaku B, vol. 13, pp. 185–333, 1968.
[51]
S. Kurita, “Chromosome number of some Japanese ferns III,” Journal of the College of Arts and Sciences, Chiba University, Natural Science Series, vol. 8, pp. 463–468, 1962.
[52]
S. M. Kawakami, M. Ito, and S. Kawakami, “Apogamous sporophyte formation in a fern Pteris multifida and its characteristics,” Journal of Plant Research, vol. 108, no. 2, pp. 181–184, 1995.
[53]
K. Mitui, “Chromosome studies on Japanese ferns,” Journal of Japanese Botany, vol. 41, pp. 61–64, 1966.
[54]
R. E. Holttum and S. K. Roy, “Cytological observations on ferns from New Guinea with descriptions of new species,” Blumea, vol. 13, pp. 129–139, 1965.
[55]
K. Mitui, “Chromosome studies on Japanese ferns,” Journal of Japanese Botany, vol. 42, pp. 105–110, 1967.
[56]
T. Nakaike, New Flora of Japan Pteridophyta, Shibundo Company, Tokyo, Japan, 1992.
[57]
N. Nakato and M. Kato, “Chromosome numbers of ten species of ferns from Guam, S. Mariana Isls.,” Acta Phytotaxonomica et Geobotanica, vol. 52, pp. 125–133, 2001.
[58]
J. Srivastava, S. A. Ranade, and P. B. Khare, “Distribution and threat status of the cytotypes of Pteris vittata L. (Pteridaceae) species complex in India,” Current Science, vol. 93, no. 1, pp. 81–85, 2007.
[59]
H. He and L. B. Zhang, “Pteris xiaoyingae, sp. nov. (sect. Pteris) from a karst cave in China based on morphological and palynological evidence,” Systematic Botany, vol. 35, no. 4, pp. 695–700, 2010.
[60]
S.-J. Lin, M. Kato, and K. Iwatsuki, “Electrophoretic variation of the apogamous Dryopteris varia group (Dryopteridaceae),” Journal of Plant Research, vol. 108, no. 4, pp. 451–456, 1995.
[61]
S.-J. Lin, M. Kato, and K. Iwatsuki, “Diploid and triploid offspring of triploid agamosporous fern Dryopteris pacifica,” The Botanical Magazine Tokyo, vol. 105, no. 3, pp. 443–452, 1992.
[62]
L. Comai, “The advantages and disadvantages of being polyploid,” Nature Reviews Genetics, vol. 6, no. 11, pp. 836–846, 2005.
[63]
R. J. Singh, Plant Cytogenetics, 2003.
[64]
Y. Watano and K. Iwatsuki, “Genetic variation in the “Japanese apogamous form” of the fern Asplenium unilaterale Lam,” The Botanical Magazine Tokyo, vol. 101, no. 3, pp. 213–222, 1988.
[65]
K. U. Kramer and P. M. McCarthy, Flora of Australia: Ferns, Gymnosperms and Allied Genera, Collingwood, 1998.
[66]
J. Jha and B. M. Sinha, “Cytomorphological variability in apogamous populations of Pteris cretica L.,” Caryologia, vol. 40, pp. 71–78, 1987.