Abstract
Interspecific hybridization had been long recognized as a widespread evolutionary process in vascular plants. In the present review, we summarize knowledge concerning studies of interspecific hybridization in bryophytes before and after the advent of molecular methods. The available data indicate that hybridization is an important evolutionary phenomenon among bryophytes. Evidence for hybridization events before the molecular era is mainly based on studies of intermediacy of parental morphology. The recent molecular marker technology has revolutionized studies of hybridization, generating new insights into the genetic and evolutionary consequences of homoploid and allopolyploid speciation. The current molecular approaches support the prevalence of allopolyploidy in bryophytes. However, we anticipate that homoploid hybridization is under-reported. Finally, we suggest some directions for future studies of hybrid speciation among bryophytes.
Downloads
References
Abbott, R., Albach, D., Ansell, S., Arntzen, J.W., Baird, S.J.E., Bierne, N., Boughman, J., Brelsford, A., Buerkle, C.A., Buggs, R., Butlin, R.K., Dieckmann, U., Eroukhmanoff, F., Grill, A., Cahan, S.H., Hermansen, J.S., Hewitt, G., Hudson, A.G., Jiggins, C., Jones, J., Keller, B., Marczewski, T., Mallet, J., Martinez-Rodriguez, P., Möst, M., Mullen, S., Nichols, R., Nolte, A.W., Parisod, C., Pfennig, K., Rice, A.M., Ritchie, M.G., Seifert, B., Smadja, C.M., Stelkens, R., Szymura, J.M., Väinöla, R., Wolf, J.B.W. & Zinner, D. (2013) Hybridization and speciation. Journal of Evolutionary Biology 26: 229–246. https://doi.org/10.1111/j.1420-9101.2012.02599.x
Abderrahman, S. (1998) DNA content of two cytotypes of Funaria hygrometrica. Korean Journal of Genetics 20: 103–108.
Abderrahman, S.A. (2004) Nuclear DNA content of haploid and diploid Physcomitruim pyriforme using DAR staining. Korean Journal of Genetics 26: 245–250.
Allen, C.E. (1930) Inheritance in a hepatic. Science 71: 197–204. https://doi.org/10.1126/science.71.1834.197
Allen, C.E. (1935) The genetics of bryophytes. The Botanical Review 1: 269–291. https://doi.org/10.1007/BF02869756
Allen, C.E. (1937) Fertility and compatibility in Sphaerocarpos. Cytologia. Fujii Jubilee 1937: 494–501. https://doi.org/10.1508/cytologia.FujiiJubilaei.494
Allen, C.E. (1945) The genetics of bryophytes II. The Botanical Review 11: 260–287. https://doi.org/10.1007/BF02861195
Allendorf, F.W., Leary, R.F., Spruell, P. & Wenburg, J.K. (2001) The problems with hybrids: setting conservation guidelines. Trends in Ecology and Evolution 16: 613–622. https://doi.org/10.1016/S0169-5347(01)02290-X
Anderson, E. (1948) Hybridization of the habitat. Evolution 2: 1–9. https://doi.org/10.1111/j.1558-5646.1948.tb02726.x
Anderson, E. & Stebbins, G.L. (1954) Hybridization as an evolutionary stimulus. Evolution 8: 378–388. https://doi.org/10.2307/2405784
Anderson, L.E. (1963) Modern species concepts: mosses. The Bryologist 46: 47–66.
Anderson, L.E. (1980) Cytology and reproductive biology of mosses. In: Taylor, R.J. & Levontin, L.E. (Eds.) The mosses of North America. California Academy of Sciences, San Francisco, California, pp. 37–76.
Anderson, L.E. & Lemmon, B.E. (1972) Cytological Studies of Natural Intergeneric Hybrids and Their Parental Species in the Moss Genera, Astomum and Weissia. Annals of the Missouri Botanical Garden 59: 382–416. https://doi.org/10.2307/2395151
Andrews, A.L. (1918) Bryological notes IV: A new hybrid in Physcomitrium. Torreya 18: 52–54.
Andrews, A.L. (1922) Hymenostomum in North America. II. The case of Astomum sullivantii. The Bryologist 25: 66–71. https://doi.org/10.2307/3238657
Andrews, A.L. (1960) Taxonomic notes. XV. The reciprocal hybrid of Ditrichum-Pleuridium. The Bryologist 63: 179–181. https://doi.org/10.2307/3241054
Andrews, A.L. & Hermann, F.J. (1959) A natural hybrid in the Ditrichaceae. The Bryologist 62: 119–122. https://doi.org/10.2307/3240030
Arnold, S.J., Avise, J.C., Ballou, J., Eldridge, J. & Flemming, D. (1991) Genetic management considerations for threatened species with a detailed analysis of the Florida panther (Felis concolor coryi). Washington, DC: USFWS.
Bainard, J.D. & Newmaster, S.G. (2010) Endopolyploidy in bryophytes: Widespread in mosses and absent in liverworts. Journal of Botany 2010: 1–7. https://doi.org/10.1155/2010/316356
Barbulescu, E.V.I., Patzak, S.D.F., Feldberg, K., Schäfer-Verwimp, A., Rycroft, D.S., Renner, M.A.M. & Heinrichs, J. (2017) Allopolyploid origin of the leafy liverwort Plagiochila britannica (Plagiochilaceae). Botanical Journal of the Linnean Society 183: 250–259. https://doi.org/10.1093/botlinnean/bow005
Bell, N.E. & Hyvönen, J. (2010) Phylogeny of the moss class Polytrichopsida (BRYOPHYTA): Generic-level structure and incongruent gene trees. Molecular Phylogenetics and Evolution 55: 381–398. https://doi.org/10.1016/j.ympev.2010.02.004
Bhatia, S., Negi, M.L. & Lakshmikumaran, M. (1996) Structural analysis of the rDNA intergenic space of Brassica nigra: evolutionary divergence of the spacers of the three diploid Brassica species. Journal of Molecular Evolution 43: 460–468. https://doi.org/10.1007/BF02337518
Bischler, H. & Boisselier-Dubayle, M.C. (1993) Variation in a polyploid, dioicous liverwort, Marchantia globosa. American Journal of Botany 80: 953–958. https://doi.org/10.1002/j.1537-2197.1993.tb15317.x
Blackstock, N. & Ashton, P.A. (2010) Genetic markers and morphometric analysis reveal past hybridization and introgression in putative Carex flava L. s.str. (Cyperaceae) hybrid populations. Plant Systematics and Evolution 287: 37–47. https://doi.org/10.1007/s00606-010-0287-0
Boisselier-Dubayle, M.C. & Bischler, H. (1998) Allopolyploidy in the thalloid liverwort Corsinia (Marchantiales). Botanica Acta 111: 490–496. https://doi.org/10.1111/j.1438-8677.1998.tb00737.x
Boisselier-Dubayle, M.C. & Bischler, H. (1999) Genetic relationships between haploid and triploid Targionia (Targioniaceae, Hepaticae). International Journal of Plant Sciences 160: 1163–1169. https://doi.org/10.1086/314199
Boisselier-Dubayle, M.C., Lambourdiere, J. & Bischler, H. (1998a) The leafy liverwort Porella baueri (Porellaceae) is an allopolyploid. Plant Systematics and Evolution 210: 175–197. https://doi.org/10.1007/BF00985667
Boisselier-Dubayle, M.C., Lambourdiere, J. & Bischler, H. (1998b) Taxa delimitation in Reboulia investigated with morphological, cytological, and isozyme markers. The Bryologist 101: 61–69. https://doi.org/10.2307/3244074
Bremer, B. (1981) A Taxonomic Revision of Schistidium (Grimmiaceae, Bryophyta) 3. Lindbergia 7: 73–90.
Buczkowska, K., Odrzykoski, I.J. & Chudzinska, E. (2004) Delimitation of some European species of Calypogeia Raddi (Jungermanniales, Hepaticae) based on cytological characters and multienzyme phenotype. Nova Hedwigia 78: 147–163. https://doi.org/10.1127/0029-5035/2004/0078-0147
Buczkowska, K., Sawicki, J., Szczecińska, M., Klama, H. & Bączkiewicz, A. (2012) Allopolyploid speciation of Calypogeia sphagnicola (Jungermanniopsida, Calypogeiaceae) based on isozyme and DNA markers. Plant Systematics and Evolution 298: 549–560. https://doi.org/10.1007/s00606-011-0565-5
Buerkle, C.A., Morris, R.J., Asmussen, M.A. & Rieseberg, L.H. (2000) The likelihood of homoploid hybrid speciation. Heridity 84: 441–451. https://doi.org/10.1046/j.1365-2540.2000.00680.x
Burgeff, H. (1943) Genetische Studien an Marchantia. Gustav Fischer, Jena, 296 pp.
Burke, J.M. & Arnold, M.L. (2001) Genetics and the fitness of hybrids. Annual Review of Genetics 35: 31–52. https://doi.org/10.1146/annurev.genet.35.102401.085719
Buryová, B. (2004) Genetic variation in two closely related species of Philonotis based on isozyme. The Bryologist 107: 316–327. https://doi.org/10.1639/0007-2745(2004)107[0316:GVITCR]2.0.CO;2
Callaghan, D.A., Masson, J. & During, H. (2020a) Physcomitrium × stevensoni D.A. Callaghan (Physcomitrium patens × P. eurystomum) (Funariaceae, Bryophyta), a new name for a rarely recorded hybrid moss. Journal of Bryology 42: 192–194. https://doi.org/10.1080/03736687.2020.1725325
Callaghan, D.A., Medina, R., Masson, J. & During, H.J. (2020) Population status and ecology of the episodic moss Physcomitrium eurystomum Sendtn. in Britain. Journal of Brylogy 42 (3): 246–257. https://doi.org/10.1080/03736687.2020.1743562
Campbell, E.O. (1971) Problems in the origin and classification of bryophytes with particular reference to liverworts. New Zealand Journal of Botany 9: 678–688. https://doi.org/10.1080/0028825X.1971.10430230
Carrión, J.S., Ros, R.M. & Guerra, J. (1993) Spore morphology in Pottia starckeana (Hedw.) C. Müll. (Pottiaceae, Musci) and its closest species. Nova Hedwigia 56: 89–112.
Chudzinska, E. & Odrzykoski, I.J. (2010) Fast identification method for an allopolyploid liverwort Pellia borealis Lorbeer (Hepaticae, Metzgeriales). Cryptogamie Bryologie 31: 207–215.
Clarke, G.C.S. (1979) Spore morphology and bryophyte systematics. In: Clarke, G.C.S. & Duckett, J.G. (Eds.) Bryophyte Systematics. Academic Press, London, pp. 231–250.
Clarke, S.M., Henry, H.M., Dodds, K.G., Jowett, T.W.D., Manley, T.R., Anderson, R.M. & Mcewan, J.C. (2014) A High Troughput Single Nucleotide Polymorphism Multiplex Assay for Parentage Assignment in New Zealand Sheep. PLoS ONE 9: e93392. https://doi.org/10.1371/journal.pone.0093392
Cox, C.J., Goffinet, B., Shaw, A.J. & Boles, S.B. (2004) Phylogenetic relationships among the mosses based on heterogeneous Bayesian analysis of multiple genes from multiple genomic compartments. Systematic Botany 29: 234–250. https://doi.org/10.1600/036364404774195458
Coyne, J.A. & Orr, H.A. (2004) Speciation. Sunderland, Massachusetts, Sinauer Associates, 545 pp.
Cronberg, N. (1989) Patterns of variation in morphological characters and isoenzymes in populations of Sphagnum capillifolium (Ehrh.) Hedw. and S. rubellum Wils. from two bogs in southern Sweden. Journal of Bryology 15: 683–696. https://doi.org/10.1179/jbr.1989.15.4.683
Cronberg, N. (1992) Reproductive Biology of Sphagnum. Lindbergia 17: 69–82.
Cronberg, N. (1996) Isozyme evidence of relationships within Sphagnum sect. Acutifolia (Sphagnaceae, Bryophyta). Plant Systematics and Evolution 203: 41–64. https://doi.org/10.1007/BF00985236
Cronberg, N. & Natcheva, R. (2002) Hybridization between the peat mosses, Sphagnum capillifolium and S. quinquefarium (Sphagnaceae, Bryophyta) as inferred by morphological characters and isozyme markers. Plant Systematics and Evolution 234: 53–70. https://doi.org/10.1007/s00606-002-0196-y
Crundwell, A.C. & Nyholm, E. (1964) The European species of the Bryum erythrocarpum complex. Transactions of the British Bryological Society 4: 597–637. https://doi.org/10.1179/006813864804812128
Crundwell, A.C. & Nyholm, E. (1974) Funaria muhlenbergii and Related European Species. Lindbergia 2: 222–229.
Crundwell, A.C. & Nyholm, E. (1977) Dicranella howei Ren. & Card. and Its Relationship to D. varia (Hedw.) Schimp. Lindbergia 4: 35–38.
Delgadillo, M.C. (1989) Astomiopsis × altivallis (Musci: Ditrichaceae), a putative interspecific hybrid in Mexico. The Bryologist 92: 225–227. https://doi.org/10.2307/3243950
Derda, G.S. & Wyatt, R. (2000) Isozyme evidence regarding the origins of three allopolyploid species of Polytrichastrum (Polytrichaceae, Bryophyta). Plant Systematics and Evolution 220: 37–53. https://doi.org/10.1007/BF00985369
Dobzhansky, T. (1937) Genetics and the origin of species. Columbia University Press, New York.
Doyle, W.T. (1960) Maternal inheritance and interspecific hybrids of the liverwort Sphaerocarpos. Nature 187: 429–430. https://doi.org/10.1038/187429a0
Ellstrand, N.C. & Elam, D.R. (1993) Population genetic consequences of small population size: implications for plant conservation. Annual Review of Ecology and Systematics 24: 217–242. https://doi.org/10.1146/annurev.es.24.110193.001245
Fiedorow, P., Odrzykoski, I., Szweykowski, J. & Szweykowska-Kulinska, Z. (2001) Phylogeny of the European species of the genus Pellia (Hepaticae; Metzgeriales) based on the molecular data from nuclear tRNA(CAA)(Leu) intergenic sequences. Gene 262: 309–315. https://doi.org/10.1016/S0378-1119(00)00523-0
Flatberg, K.I. (1988) Sphagnum angustifolium (Russ.) C. Jens. × S. pulchrum (Braithw.) Warnst., a gametophyte hybrid. Lindbergia 14: 4–7.
Flatberg, K.I. (2005) Taxonomy, Geography and Possible Origin of Sphagnum inexspectatum (Sect. Subsecunda) sp. nov. Lindbergia 30: 59–78.
Flatberg, K.I. & Frisvoll, A.A. (1984) Sphagnum arcticum sp. nov. The Bryologist 87: 143–148. https://doi.org/10.2307/3243120
Flatberg, K.I., Thingsgaard, K. & Såstad, S.M. (2006) Interploidal gene flow and introgression in bryophytes: Sphagnum girgensohnii × Sphagnum russowii, a case of spontaneous neotriploidy. Journal of Bryology 28: 27–37. https://doi.org/10.1179/174328206X90459
Frahm, J. & Ho, B. (2010) Discovery of a natural hybrid between Bruchia vogesiaca Schwagr. and Trematodon ambiguus (Hedw.) Hornsch (Musci, Bruchiaceae). Cryptogamie Bryologie 31: 95–99.
Freeland, J.R., Kirk, H. & Petersen, S.D. (2011) Molecular Ecology. 2nd ed. Wiley-Blackwell. 464 pp. https://doi.org/10.1002/9780470979365
Frisvoll, A.A. (1978) The genus Tetraplodon in Norway, a taxonomic revision. Lindbergia 4: 225–246.
Fritsch, R. (1982) Index to plant chromosome numbers-Bryophyta. Regnum Vegetabile 108: 1–140.
Gallego, M.T., Cano, M.J., Ros, R.M. & Guerra, J. (2002) New Taxonomic Data on a Circum-Tethyan Group of Syntrichia (Pottiaceae, Bryophyta): the S. caninervis Complex. Systematic Botany 27: 643–653.
Gibson, D.J. (2015) Methods in comparative plant population ecology. 2nd ed. Oxford University Press, 320 pp.
Grant, V. (1975) Genetics of flowering plants. 2nd edition. Columbia University Press, New York, USA.
Grant, V. (1981) Plant Speciation. New York: Columbia University Press. New York, USA, 563 pp.
Gross, B.L. & Rieseberg, L.H. (2005) The ecological genetics of homoploid hybrid speciation. Journal of Heredity 96: 241–252. https://doi.org/10.1093/jhered/esi026
Guerra, J., Ros, R.M. & Cano, M.J. (1994) Pterygoneurum subsessile (Brid.) Jur. var. kieneri Hab. (Musci, Pottiaceae), a putative hybrid. Nova Hedwigia 58: 507–510.
Györffy, I. (1907) Ueber die vergleichenden anatomischen Verhältnisse von Physcomitrella patens (Hedw.) Br. et Sch., Physcomitrium pyriforme (L.) Brid., Physcomitrium sphaericum (Ludw.) Brid., und Physcomitrella Hampei Limpr. [Physcomitrella patens (Hedw.) × Physcomitrium sphaericum (Ludw.), resp.: Physcomitrella patens (Hedw.) × Physcomitrium pyriforme (L.)]. Hedwigia 47: 1–59.
Harrison, R.G. & Larson, E.L. (2014) Hybridization, introgression, and the nature of species boundaries. Journal of Heredity 105: 795–809. https://doi.org/10.1093/jhered/esu033
Hedderson, T.A. (1986) A naturally occurring moss hybrid between Orthotrichum gymnostomum and O. obtusifolium from Newfoundland, Canada. The Bryologist 89: 165–167. https://doi.org/10.2307/3242759
Hedenäs, L. (2015) Tortella rigens (Bryophyta, Pottiaceae): relationships, regional variation, and conservation aspects. Plant Systematics and Evolution 301: 1361–1375. https://doi.org/10.1007/s00606-014-1159-9
Hegarty, M.J. & Hiscock, S.J. (2005) Hybrid speciation in plants: new insights from molecular studies. New Phytologist 165: 411–423. https://doi.org/10.1111/j.1469-8137.2004.01253.x
Heinrichs, J., Kreier, H.P., Feldberg, K., Schmidt, A.R., Zhu, R.L., Shaw, B., Shaw, A.J. & Wissemann, V. (2011) Formalizing morphologically cryptic biological entities: New insights from DNA taxonomy, hybridization, and biogeography in the leafy liverwort Porella platyphylla (Jungermanniopsida, Porellales). American Journal of Botany 98: 1252–1262. https://doi.org/10.3732/ajb.1100115
Hernández-Maqueda, R., Quandt, D. & Muñoz, J. (2008) Testing reticulation and adaptive convergence in the Grimmiaceae (Bryophyta). Taxon 57: 500–510.
Hofmann, H. (1998) A monograph of the genus Homalothecium (Brachytheciaceae, Musci). Lindbergia 23: 119–159.
Holmen, K. & Scotter, G.W. (1971) Mosses of the Reindeer Preserve, Northwest Territories, Canada. Lindbergia 1: 34–56.
Hugonnot, V. (2017) Comparative investigations of niche, growth rates and reproduction between the native moss Campylopus pilifer and the invasive C. introflexus. Journal of Bryology 39: 79–84. https://doi.org/10.1080/03736687.2016.1210871
Hylander, K. (1999) Intermediates between the pleurocarpous mosses Dichelyma falcatum (Hedw.) Myr. And D. capillaceum (Dicks.) Myr., possible of hybrid origin. Lindbergia 24: 59–64.
Ignatov, M.S. & Milyutina, I.A. (2011) Intrafamilial hybridization in mosses? an enigmatic case in the genus Podperaea (Hypnales, Bryophyta). Arctoa 20: 107–118. https://doi.org/10.15298/arctoa.20.08
Ignatov, M.S., Kuznetsova, O.I. & Ignatova, E.A. (2019) Hybridization in mosses and how remote it could be. Biology Bulletin Reviews 9: 267–273. https://doi.org/10.1134/S207908641903006X
Ireland, R. (1987) Scanning electron microscope study of the spores of the North American species of Plagiothecium. Memoirs of the New York Botanical Garden 45: 95–110.
Jankowiak, K., Rybarczyk, A., Wyatt, R., Odrzykoski, I., Pacak, A. & Szweykowska-Kulinska, Z. (2005) Organellar Inheritance in the Allopolyploid Moss Rhizomnium pseudopunctatum. Taxon 54: 383–388. https://doi.org/10.2307/25065367
Jankowiak, K. & Szweykowska-Kulinska, Z. (2004) Organellar inheritance in the allopolyploid liverwort species Porella baueri (Porellaceae): Reconstructing historical events using DNA analysis data. In: Goffinet, B., Hollowell, V. & Magill, R. (Eds.) Molecular Systematics of Bryophytes, Missouri Botanical Garden Press, St Louis, pp. 404–414.
Jankowiak-Siuda, K., Pacak, A., Odrzykoski, I., Wyatt, R. & Szweykowska-Kulinska, Z. (2008) Organellar inheritance in the allopolyploid moss Plagiomnium curvatulum. Taxon 57: 145–152.
Jin, Y., Liu, S., Yuan, Z., Yang, Y., Tan, S. & Liu, Z. (2016) Catfish genomic studies: progress and perspectives. In: MacKenzie, S. & Jentoft, S. (Eds.) Genomics in Aquaculture. Elsevier Inc., pp. 73–104. https://doi.org/10.1016/B978-0-12-801418-9.00004-4
Karlin, E.F. (2014) Subgenome analysis of two southern hemisphere allotriploid species in Sphagnum (Sphagnaceae). Journal of Bryology 36: 165–179. https://doi.org/10.1179/1743282014Y.0000000098
Karlin, E.F., Boles, S.B., Ricca, M., Temsch, E., Greilhuber, J. & Shaw, A.J. (2009) Three-genome mosses: complex double allopolyploid origins for triploid gametophytes in Sphagnum. Molecular Ecology 18: 1439–1454. https://doi.org/10.1111/j.1365-294X.2009.04113.x
Karlin, E.F., Buck, W.R., Seppelt, R.D., Boles, S.B. & Shaw, A.J. (2013) The double allopolyploid Sphagnum × falcatulum (Sphagnaceae) in Tierra del Fuego, a Holantarctic perspective. Journal of Bryology 35: 157–172. https://doi.org/10.1179/1743282013Y.0000000066
Karlin, E.F., Gardner, G. P., Lukshis, K., Boles, S. & Shaw, A.J. (2010) Allopolyploidy in Sphagnum mendocinum and S. papillosum (Sphagnaceae). The Bryologist 113: 114–119. https://doi.org/10.1639/0007-2745-113.1.114
Karlin, E.F. & Smouse, P. (2017) Allo-allo-triploid Sphagnum × falcatulum: single individuals contain most of the Holantarctic diversity for ancestrally indicative markers. Annals of Botany 120: 221–231. https://doi.org/10.1093/aob/mcw269
Karlin, E.F., Temsch, E.M., Bizuru, E., Marino, J., Boles, S.B., Devos, N. & Shaw, A.J. (2014) Invisible in plain sight: recurrent double allopolyploidy in the African Sphagnum × planifolium (Sphagnaceae). The Bryologist 117: 187–201. https://doi.org/10.1639/0007-2745-117.2.187
Khanna, K.R. (1960) Studies in natural hybridization in the genus Weissia. The Bryologist 63: 1–16. https://doi.org/10.2307/3241166
Köckinger, H. & Hedenäs, L. (2017) A farewell to Tortella bambergeri (Pottiaceae) as understood over the last decades. Journal of Bryology 39: 213–225. https://doi.org/10.1080/03736687.2017.1307313
Košnar, J. & Kolář, F. (2009) A taxonomic study of selected European taxa of the Tortula muralis (Pottiaceae, Musci) complex: variation in morphology and ploidy level. Preslia 81: 399–421.
Košnar, M.J., Miroslava, H., Kolář, F., Koutecký, P. & Kučera, J. (2012) A case study of intragenomic ITS variation in bryophytes: Assessment of gene flow and role of polyploidy in the origin of European taxa of the Tortula muralis (Musci: Pottiaceae) complex. Taxon 61: 709–720. https://doi.org/10.1002/tax.614001
Kugita, M., Kaneko, A., Yamamoto, Y., Takeya, Y., Matsumoto, T. & Yoshinaga, K. (2003) The complete nucleotide sequence of the hornwort (Anthoceros formosae) chloroplast genome: insight into the earliest land plants. Nucleic Acids Research 31: 716–721. https://doi.org/10.1093/nar/gkg155
Kyrkjeeide, M.O., Hassel, K., Aguero, B., Temsch, E.M., Afonina, O.M., Shaw, A.J., Stenøien, H.K. & Flatberg, K.I. (2019). Sphagnum × lydiae, the first allotriploid peatmoss in the northern hemisphere. The Bryologist 122: 38–61. https://doi.org/10.1639/0007-2745-122.1.038
Lal, M. (1984) The Experimental Biology of Bryophytes. Academic Press, London, pp. 97–115.
Levin, D.A., Francisco-Ortega, J. & Jansen, R.K. (1996) Hybridization and the extinction of rare species. Conservation Biology 10: 10–16. https://doi.org/10.1046/j.1523-1739.1996.10010010.x
Linde, A-M., Sawangproh, W., Cronberg, N., Szövényi, P. & Lagercrantz, U. (2020) Evolutionary history of the Marchantia polymorpha complex. Frontiers in Plant Science. https://doi.org/10.3389/fpls.2020.00829
Liu, Y., Moskwa, N.L. & Goffinet, B. (2012) Development of eight mitochondrial markers for Funariaceae (Musci) and their amplification success in other mosses. American Journal of Botany 99: 62–65. https://doi.org/10.3732/ajb.1100402
Lobachevs’ka, O.V., Demkiv, O.T. & Ripets’kyi, R.T. (1986) Cytophotometric determination of nuclear DNA in the polyploid complex Tortula muralis Hedw. Ukrayins’kyi Botanichnyi Zhurnal 43: 23–26.
Lobachevs’ka, O.V. & Ulychna, K.O. (1994) Peculiarities of hybrid population of moss Tortula canescens Mont. times T. muralis Hedw. from vicinities of Nikolayev (Lviv district). Ukrayins’kyi Botanichnyi Zhurnal 51: 84–91.
Lysak, M.A., Fransz, P.F., Ali, H.B.M. & Schubert, I. (2001) Chromosome painting in Arabidopsis thaliana. Plant Journal 28: 689–697. https://doi.org/10.1046/j.1365-313x.2001.01194.x
Maki, M. & Murata, J. (2001) Allozyme analysis of the hybrid origin of Arisaema ehimense (Araceae). Heredity 86: 87–93. https://doi.org/10.1046/j.1365-2540.2001.00813.x
Mallet, J. (2007) Hybrid speciation. Nature 446: 279–283. https://doi.org/10.1038/nature05706
Mayr, E. (1942) Systematics and the origin of species. Columbia University Press, New York, 334 pp.
McDaniel, S.F., von Stackelberg, M., Richardt, S., Quatrano, R.S., Reski, R. & Rensing, S.A. (2010) The speciation history of the Physcomitrium—Physcomitrella species complex. Evolution 64: 217–231. https://doi.org/10.1111/j.1558-5646.2009.00797.x
McDaniel, S.F., Willis, J.H. & Shaw, A.J. (2007) Linkage map reveals a complex basis for segregation distortion in an interpopulation cross in the moss Ceratodon purpureus. Genetics 176: 2489–2500. https://doi.org/10.1534/genetics.107.075424
McDaniel, S.F., Willis, J.H. & Shaw, A.J. (2008) The Genetic Basis of Developmental Abnormalities in Interpopulation Hybrids of the Moss Ceratodon purpureus. Genetics 179: 1425–1435. https://doi.org/10.1534/genetics.107.086314
McIntosh, T.T. (1989) Bryophyte Records from the Semiarid Steppe of Northwestern North America, including Four Species New to North America. The Bryologist 92: 356–362. https://doi.org/10.2307/3243404
Meleshko, O., Stenøien, H.K., Speed, J.D.M., Flatberg, K., Kyrkjeeide, M.O. & Hassel, K. (2018) Is interspecific gene flow and speciation in peatmosses (Sphagnum) constrained by phylogenetic relationship and life-history traits? Lindbergia 41: linbg.01107, 2018. https://doi.org/10.25227/linbg.01107
Natcheva, R. & Cronberg, N. (2004) What do we know about hybridization among bryophytes in nature? Canadian Journal of Botany 82: 1687–1704. https://doi.org/10.1139/b04-139
Natcheva, R. & Cronberg, N. (2007a) Maternal transmission of cytoplasmic DNA in interspecific hybrids of peat mosses, Sphagnum (Bryophyta). Journal of Evolutionary Biology 20: 1613–1616. https://doi.org/10.1111/j.1420-9101.2007.01341.x
Natcheva, R. & Cronberg, N. (2007b) Recombination and introgression of nuclear and chloroplast genomes between the peat mosses, Sphagnum capillifolium and Sphagnum quinquefarium. Molecular Ecology 16: 811–818. https://doi.org/10.1111/j.1365-294X.2006.03163.x
Newton, M.E. (1983) Cytology of the Hepaticae and Anthocerotae. In: Schuster, R.M. (Ed.) New Manual of Bryology, Vol. 1. The Hattori Botanical Laboratory, Nichinan, p. 1295.
Newton, M.E. (1988) Chromosomes as indicators of bryophyte reproductive performance. Botanical Journal of the Linnean Society 98: 269–275. https://doi.org/10.1111/j.1095-8339.1988.tb02430.x
Nickrent, D.L., Parkinson, C.L., Palmer, J.D. & Duff, R.J. (2000) Multigene Phylogeny of Land Plants with Special Reference to Bryophytes and the Earliest Land Plants. Molecular Biology and Evolution 17: 1885–1895. https://doi.org/10.1093/oxfordjournals.molbev.a026290
Nicholson, W.E. (1905) Notes on two forms of hybrid Weisia. Revue Bryologique 32: 19–25.
Nicholson, W.E. (1906) Weisia crispa Mitt. × W. microstoma C.M. Revue Bryologique 33: 1–2.
Nicholson, W.E. (1910) A new hybrid moss. Revue Bryologique 37: 23–24.
Nicholson, W.E. (1931) Hybridity among the bryophytes. Revue Bryologique 4: 138–140.
Nieto-Lugilde, M. (2019) The moss genus Ceratodon: Studies of Evolutionary Biology and Taxonomy in southern Europe. PhD thesis, University of Murcia, Spain.
Nieto-Lugilde, M., Werner, O., McDaniel, S.F. & Ros-Espín, R.M. (2018) Environmental variation obscures species diversity in southern European populations of the moss genus Ceratodon. Taxon 67: 673–692. https://doi.org/10.12705/674.1
Norrell, T.E., Jones, K.S., Payton, A.C. & McDaniel, S.F. (2014) Meiotic sex ratio variation in natural populations of Ceratodon purpureus (Ditrichaceae). American Journal of Botany 101: 1572–1576. https://doi.org/10.3732/ajb.1400156
Nyholm, E. (1954) Illustrated Moss Flora of Fennoscandia. II. Musci. Fasc. I. Lund. Gleerup
Nyholm, E. (1958) Illustrated Moss Flora of Fennoscandia. II. Musci. Fasc. III. Lund. Gleerup.
Odrzykoski, I.J., Chudzińska, E. & Szweykowski, J. (1996) The hybrid origin of the polyploidy liverwort Pellia borealis. Genetica 98: 75–86. https://doi.org/10.1007/BF00120221
Orzechowska, M., Karcz, J. & Małuskyńska, J. (2006) Comparative analysis of the structure of the allopolyploid liverwort Pellia borealis and ancestral taxa. Biodiversity Research and Conservation 1–2: 54–56.
Orzechowska, M., Siwinska, D. & Maluszynska, J. (2010) Molecular cytogenetic analyses of haploid and allopolyploid Pellia species. Journal of Bryology 32: 113–121. https://doi.org/10.1179/037366810X12578498136075
Pacak, A. & Szweykowska-Kulinska, Z. (2003) Organellar inheritance in liverworts: An example of Pellia borealis. Journal of Molecular Evolution 56: 11–17. https://doi.org/10.1007/s00239-002-2375-4
Pardo, D.C., Terracciano, S., Giordano, S. & Spagnuolo, V. (2014) Molecular Markers Based on PCR Methods: A Guideline for Mosses. Cryptogamie, Bryologie 35: 229–246. https://doi.org/10.7872/cryb.v35.iss3.2014.229
Parker, P.G., Snow, A.A., Schug, M.D., Booton, G.C. & Fuerst, P.A. (1998) What molecules can tell us about populations: choosing and using a molecular marker. Ecology 79: 361–382. https://doi.org/10.2307/176939
Pereira, M.R., Ledent, A., Mardulyn, P., Zartman, C.E. & Vanderpoorten, A. (2019) Maintenance of genetic and morphological identity in two sibling Syrrhopodon species (Calymperaceae, Bryopsida) despite extensive introgression. Journal of Systematics and Evolution 57: 395–403. https://doi.org/10.1111/jse.12502
Persson, H. (1954) Mosses of Alaska-Yukon. The Bryologist 57: 189–217. https://doi.org/10.2307/3240083
Petit, C., Bretagnolle, F. & Felber, F. (1999) Evolutionary consequences of diploid-polyploid hybrid zones in wild species. Trends in Ecology & Evolution 14: 306–311. https://doi.org/10.1016/S0169-5347(99)01608-0
Pettet, A. (1964) Hybrid sporophytes in the Funariaceae. I. hybrid sporophytes on Physcomitrella patens (Hedw.) B. & S. and Physcomitrium sphaericum (Schkuhr) Brid. In Britain. Transactions of the British Bryological Society 4: 642–648. https://doi.org/10.1179/006813864804812164
Pócs, T., Sabovljevic, M., Puche, F., Moragues, J.G.S., Gimeno, C. & Kürschner, H. (2004) Crossidium laxefilamentosum Frey & Kürschner (Bryopsida: Pottiaceae), new to Europe and to North Africa. Journal of Bryology 26: 113–124. https://doi.org/10.1179/037366804225021074
Proctor, V.W. (1972) The genus Riella in North and South America: Distribution, culture and reproductive isolation. The Bryologist 75: 281–289. https://doi.org/10.2307/3241465
Proskauer, J. (1969) Studies on Anthocerotales. VIII. Phytomorphology 19: 52–66.
Reese, W.D. & Lemmon, B.E. (1965) A natural hybrid between Weissia and Astomum and notes on the nomenclature of the North American species of Astomum. The Bryologist 68: 277–283. https://doi.org/10.2307/3240597
Rhymer, J.M. & Simberloff, D. (1996) Extinction by hybridization and introgression. Annual Review of Ecology and Systematics 27: 83–109. https://doi.org/10.1146/annurev.ecolsys.27.1.83
Ricca, M., Beecher, F.W., Boles, S.B., Temsch, E., Greilhuber, J., Karlin, E.F. & Shaw, A.J. (2008) Cytotype variation and allopolyploidy in North American species of the Sphagnum subsecundum complex. American Journal of Botany 95: 1606–1620. https://doi.org/10.3732/ajb.0800148
Ricca, M. & Shaw, A.J. (2010) Allopolyploidy and homoploid hybridization in the Sphagnum subsecundum complex (Sphagnaceae: Bryophyta). Biological Journal of the Linnean Society 99: 135–151. https://doi.org/10.1111/j.1095-8312.2009.01340.x
Ricca, M., Szövényi, P., Temsch, E.M., Johnson, M.G. & Shaw, A.J. (2011) Interploidal hybridization and mating patterns in the Sphagnum subsecundum complex. Molecular Ecology 20: 3202–3218. https://doi.org/10.1111/j.1365-294X.2011.05170.x
Rieseberg, L.H. (1991) Homoploid reticulate evolution in Helianthus (Asteraceae) evidence from ribosomal genes. American Journal of Botany 78: 1218–1237. https://doi.org/10.1002/j.1537-2197.1991.tb11415.x
Rieseberg, L.H. (1995) The role of hybridization in evolution: old wine in new skins. American Journal of Botany 82: 944–953. https://doi.org/10.1002/j.1537-2197.1995.tb15711.x
Rieseberg, L.H. (1997) Hybrid origins of plant species. Annual Review of Ecology and Systematics 28: 359–389. https://doi.org/10.1146/annurev.ecolsys.28.1.359
Rieseberg, L.H., Gerber, D. (1995) Hybridization in the Catalina mahogany: RAPD evidence. Conservation Biology 9: 199–203. https://doi.org/10.1046/j.1523-1739.1995.09010199.x
Rieseberg, L.H., Whitton, J. & Linder, C.R. (1996) Molecular marker incongruence in plant hybrid zones and phylogenetic trees. Acta Botanica Neerlandica 45: 143–262. https://doi.org/10.1111/j.1438-8677.1996.tb00515.x
Risse, S. (1985) Pollia intermedia (Turn.) Fürnr. with rhizoidal tubers. Journal of Bryology 13: 523–526. https://doi.org/10.1179/jbr.1985.13.4.523
Roberts, H.F. (1929) Plant Hybridization before Mendel. Princeton, New Jersey: Princeton University Press. https://doi.org/10.5962/bhl.title.4517
Ros, R.M., Guerra, J. & Cano, M.J. (1994) Pottia × andalusica (Musci: Pottiaceae), un híbrido interespecífico en Pottieae. Cryptogamie, Bryologie-Lichénologie 15: 199–204.
Ros, R.M., Guerra, J., Carrion, J.S. & Cano, M.J. (1996) A new point of view on the taxonomy of Pottia starckeana agg. (Musci, Pottiaceae). Plant Systematics and Evolution 199: 153–165. https://doi.org/10.1007/BF00984902
Rosengren, F. & Cronberg, N. (2015) Selective spore germination on shoots of Homalothecium lutescens, a moss with dwarf males. Biological Letters 11: 20150427 https://doi.org/10.1098/rsbl.2015.0427
Rusek, J., Ayan, G.B., Turko, P., Telienbach, C., Giessler, S., Spaak, P. & Wolinska, J. (2015) New possibilities arise for studies of hybridization: SNP-based markers for the multi-species Daphnia longispina complex derived from transcriptome data. Journal of Plankton Research 37: 626–635. https://doi.org/10.1093/plankt/fbv028
Rushing, A.E. & Snider, J.A. (1985) A natural hybrid between Bruchia microspora Nog., and Trematodon longicollis Michx. Monographs in systematics botany from the Missouri botanical garden 11: 121–132.
Sawangproh, W., Hedenäs, L., Lang, A.S., Hansson, B. & Cronberg, N. (2020a) Gene transfer across species boundaries in bryophytes: evidence from major life cycle stages in Homalothecium lutescens and H. sericeum. Annals of Botany 125: 565–579. https://doi.org/10.1093/aob/mcz209
Sawangproh, W., Lang, A.S., Hedenäs, L. & Cronberg, N. (2020b) Morphological characters and SNP markers suggest hybridization and introgression in sympatric populations of the pleurocarpous mosses Homalothecium lutescens and H. sericeum. Organisms Diversity & Evolution. https://doi.org/10.1007/s13127-020-00456-x
Såstad, S.M. (2005) Patterns and mechanisms of polyploid speciation in bryophytes. Regnum Vegetabile 143: 317– 334.
Såstad, S.M., Flatberg, K.I. & Cronberg, N. (1999) Electrophoretic evidence supporting a theory of allopolyploid origin of the peatmoss Sphagnum jensenii. Nordic Journal of Botany 19: 355–362. https://doi.org/10.1111/j.1756-1051.1999.tb01127.x
Såstad, S. M., Flatberg, K.I. & Hanssen, L. (2000) Origin, taxonomy and population structure of the allopolyploid peat moss Sphagnum majus. Plant Systematics and Evolution 225: 73–84. https://doi.org/10.1007/BF00985459
Såstad, S.M., Stenøien, H.K., Flatberg, K.I. & Bakken, S. (2001) The narrow endemic Sphagnum troendelagicum is an allopolyploid derivative of the widespread S. balticum and S. tenellum. Systematic Botany 26: 66–74.
Schmickl, R., Marburger, S., Bray, S. & Yant, L. (2017) Hybrids and horizontal transfer: introgression allows adaptive allele discovery. Journal of Experimental Botany 68: 5453–5470. https://doi.org/10.1093/jxb/erx297
Schuster, R.M. (1966) The Hepaticae and Anthocerotae of North America East of the Hundreth Meridian, vol. 1. Columbia University Press, New York.
Schuster, R.M. (1988) The aims and achievements of bryophyte taxonomists. Botanical Journal of the Linnean Society 98: 185–202. https://doi.org/10.1111/j.1095-8339.1988.tb02423.x
Shaw, A.J. (1994) Systematics of Mielichhoferia (Bryaceae: Musci). III. Hybridization between M. elongate and M. mielichhoferiana. American Journal of Botany 81: 782–790. https://doi.org/10.1002/j.1537-2197.1994.tb15515.x
Shaw, A.J. (1998) Genetic analysis of a hybrid zone in Mielichhoferia (Musci). In: Bates, J.W., Ashton, N.W. & Duckett, J.G. (Eds.) Bryology for the twenty-first century. Maney Publishing and the British Bryological Society, Leeds, pp. 161–174. https://doi.org/10.1201/9781315138626-12
Shaw, A.J. (2009) Bryophyte species and speciation. In: Goffinet, B. & Shaw, A.J. (Eds.) Bryophyte Biology, Cambridge: Cambridge University Press, pp. 445–485. https://doi.org/10.1017/CBO9780511754807.012
Shaw, A.J. & Bartow, S.M. (1992) Genetic Structure and Phenotypic Plasticity in Proximate Populations of the Moss, Funaria hygrometrica. Systematic Botany 17: 257–271. https://doi.org/10.2307/2419521
Shaw, A.J., Boles, S. & Shaw, B. (2008a) A phylogenetic delimitation of the “Sphagnum subsecundum complex” (Sphagnaceae, Bryophyta). American Journal of Botany 95: 731–744. https://doi.org/10.3732/ajb.0800048
Shaw, A.J., Cao, T., Wang, L.-S., Flatberg, K.I., Flatberg, B., Shaw, B., Zhou, P., Boles, S.B. & Terraccino, S. (2008c) Genetic variation in three Chinese peat mosses (Sphagnum) based on microsatellite markers, with primer information and analysis of ascertainment bias. The Bryologist 111: 271–281. https://doi.org/10.1639/0007-2745(2008)111[271:GVITCP]2.0.CO;2
Shaw, A.J., Cox, C.J., Boles, S.B. & Hoot, S.B. (2005) Phylogeny, species delimitation, and recombination in Sphagnum section Acutifolia. Systematic Botany 30: 16–33. https://doi.org/10.1600/0363644053661823
Shaw, A.J., Flatberg, K.I., Szövényi, P., Ricca, M., Johnson, M.G., Stenøien, H.K. & Shaw, B. (2012a) Systematics of the Sphagnum fimbriatum complex: phylogenetic relationships, morphological variation, and allopolyploidy. Systematic Botany 63: 351–364.
Shaw, A.J. & Goffinet, B. (2000) Molecular Evidence of Reticulate Evolution in the Peatmosses (Sphagnum), including S. ehyalinum sp. nov. The Bryologist 103: 357–374. https://doi.org/10.1639/0007-2745(2000)103[0357:MEOREI]2.0.CO;2
Shaw, A.J., Pokorny, L., Shaw, B., Ricca, M., Boles, S. & Szövényi, P. (2008b) Genetic structure and genealogy in the Sphagnum subsecundum complex (Sphagnaceae: Bryophyta). Molecular Phylogenetics and Evolution 49: 304–317. https://doi.org/10.1016/j.ympev.2008.06.009
Shaw, A J., Shaw, B. & Johnson, M.G. (2013) Origins, genetic structure and systematics of the narrow endemic peatmosses (Sphagnum): S. guwassanense and S. trieriporum (Sphagnaceae). American Journal of Botany 100: 1202–1220. https://doi.org/10.3732/ajb.1200630
Shaw, A J., Shaw, B. Ricca, M. & Flatberg, K.I. (2012b) A phylogenetic monograph of the Sphagnum subsecundum complex (Sphagnaceae) in eastern North America. The Bryologist 115: 128–152. https://doi.org/10.1639/0007-2745-115.1.128
Shaw, A.J., Shaw, B., Stenøien, H.K., Golinski, G.K., Hassel, K. & Flatberg, K.I. (2015) Pleistocene survival, regional genetic structure and interspecific gene flow among three northern peat-mosses: Sphagnum inexspectatum, S. orientale and S. miyabeanum. Journal of Biogeography 42: 364–376. https://doi.org/10.1111/jbi.12399
Smith, A.J.E. (1978) Cytogenetics, biosystematics and evolution in Bryophyta. Advances in Botanical Research 6: 195–276. https://doi.org/10.1016/S0065-2296(08)60331-6
Soltis, P.S. & Soltis. D.E. (2009) The role of hybridization in plant speciation. Annual Review of Plant Biology 60: 561–588. https://doi.org/10.1146/annurev.arplant.043008.092039
Stace, C.A. (1987) Hybridization and the plant species. In: Urbanska, K.M. (Ed.) Differentiation patterns in higher plants. Academic Press, New York. 115–127 pp.
Stark, L.R. (1987) A taxonomic monograph of Forsstroemia Lindb. (Bryopsida: Leptodontaceae). Journal of the Hattori Botanical Laboratory 63: 133–218.
Stebbins, G.L. (1950) Variation and Evolution in Plants. New York: Columbia University Press. https://doi.org/10.7312/steb94536
Stech, M. & Quandt, D. (2010) 20,000 species and five key markers: the status of molecular bryophyte phylogenetics. Phytotaxa 9: 196–228. https://doi.org/10.11646/phytotaxa.9.1.11
Stenøien, H.K. & Flatberg, K.I. (2000) Genetic variability in the rare Norwegian peat moss Sphagnum troendelagicum. The Bryologist 103: 794–801. https://doi.org/10.1639/0007-2745(2000)103[0794:GVITRN]2.0.CO;2
Stenøien, H.K., Shaw, A.J., Stengrundet, K. & Flatberg, K.I. (2011) The narrow endemic Norwegian peat moss Sphagnum troendelagicum originated before the last glacial maximum. Heredity 106: 370–382. https://doi.org/10.1038/hdy.2010.96
Sukkharak, P., Gradstein, S.R. & Stech, M. (2011) Phylogeny, taxon circumscriptions, and character evolution in the core Ptychanthoideae (Lejeuneaceae, Marchantiophyta). Taxon 60: 1607–1622. https://doi.org/10.1002/tax.606006
Sytsma, K.J. (1990) DNA and morphology: inference of plant phylogeny. Trends in Ecology & Evolution 5: 104–110. https://doi.org/10.1016/0169-5347(90)90163-8
Todesco, M., Pascual, M.A., Owens, G.L., Ostevik, K.L., Moyers, B.T., Hübner, S., Heredia, S.M., Hahn, M.A., Caseys, C., Bock, D.G. & Rieseberg, L.H. (2016) Hybridization and extinction. Evolutionary Applications 9: 892–908. https://doi.org/10.1111/eva.12367
Ulychna, K.O. (1977) Hybrid sporogones in Phascum-Cuspidatum Pottiaceae Musci. Ukrayins’kyi Botanichnyi Zhurnal 34: 155–158.
Vanderpoorten, A. & Goffinet, B. (2009) Introduction to Bryophytes. Cambridge University Press, UK. https://doi.org/10.1017/CBO9780511626838
Vanderpoorten, A., Hedenäs, L. & Jacquemart, A.L. (2003) Differentiation in DNA fingerprinting and morphology among species of the pleurocarpous moss genus Phytidiadelphus (Hylocomiaceae). Taxon 52: 229–236. https://doi.org/10.2307/3647391
Vanderpoorten, A. & Shaw, A.J. (2010) The application of molecular data to the phylogenetic delimitation of species in bryophytes: A note of caution. Phytotaxa 9: 229–237. https://doi.org/10.11646/phytotaxa.9.1.12
van der Velde, M. & Bijlsma, R. (2001) Genetic Evidence for the Allodiploid Origin of the Moss Species Polytrichum longisetum. Plant Biology 3: 379–385. https://doi.org/10.1055/s-2001-16453
van der Velde, M. & Bijlsma, R. (2004) Hybridization and asymmetric reproductive isolation between the closely related bryophyte taxa Polytrichum commune and P. uliginosum. Molecular Ecology 13: 1447–1454. https://doi.org/10.1111/j.1365-294X.2004.02154.x
Vicente, M.C. de & Tanksley, S.D. (1993) QTL analysis of transgressive segregation in an interspecific tomato cross. Genetics 134: 585–596. https://doi.org/10.1093/genetics/134.2.585
Vilnet, A., Konstantinova, N. & Troitsky, A. (2012) Molecular phylogenetic data on reticulate evolution in the genus Barbilophozia Loske (Anastrophyllaceae, Marchantiophyta) and evidence of non-concerted evolution of rDNA in Barbilophozia rubescens allopolyploid. Phytotaxa 49: 6–22. https://doi.org/10.11646/phytotaxa.49.1.2
Vogelmayer, H. (1998) Genome size analyses in mosses (Musci) and downy mildews (Peronosporales). PhD thesis, University of Vienna, Vienna.
von Wettstein, F. (1924) Morphologie und physiologie des formwechsels der moose auf genetischer grundlage I. Zeitschrift für inductive Abstammungs-und Vererbungeslehre 33: 1–236. https://doi.org/10.1007/BF01762372
von Wettstein, F. (1928) Morphology and physiology of moss variation on a genetic basis. II. E Baur Bibliotheca Genetica X, 216 pp.
von Wettstein F. (1932) Genetik. In: Verdoorn, F. (Ed.) Manual of bryology. Hague: Martinus Nijhoff, pp. 233–272.
Vries, A. de, Bramer, J.P.J., Zanten, B.O. van, Hofman, A. & Bijlsma, R. (1989) Allozyme Variation in Populations of Four Racopilum Species, including the Polyploid R. tomentosum. Lindbergia 15: 47–59.
Wagenitz, G. (2003) Wörterbuch der Botanik: Die Termini in ihrem historischen Zusammenhang. Spektrum Akademischer Verlag, Heidelberg.
Welsh, J. & McClelland, M. (1990) Fingerprinting genomes using PCR with arbitrary primers. Nucleic Acid Research 18: 7213–7218. https://doi.org/10.1093/nar/18.24.7213
Werner, O., Köckinger, H., Magdy, M. & Ros, R.M. (2014) On the systematic position of Tortella arctica and Trichostomum arcticum (Bryophyta, Pottiaceae). Nova Hedwigia 98: 273–293. https://doi.org/10.1127/0029-5035/2014/0175
Whitham, T.G., Morrow, P.A. & Potts, B.M. (1994) Plant hybrid zones as center for biodiversity: the herbivore community of two endemic Tasmanian eucalypts; Oecologia 97: 481–490. https://doi.org/10.1007/BF00325886
Williams, C. (1966) A natural hybrid in the genus Weissia. The Bryologist 69: 361–365. https://doi.org/10.2307/3240835
Wissemann, V. (2007) Plant evolution by means of hybridization. Systematic and Biodiversity 5: 243–253.
https://doi.org/10.1017/S1477200007002381
Wojnicka-Poltorak, A., Chudzinska, E. & Prus-Glowacki, W. (1997) Inter-and intraspecific serological relationships in Pellia epiphylla complex. Acta Societatis Botanicorum Poloniae 66: 365–370. https://doi.org/10.5586/asbp.1997.045
Wyatt, R. (1994) Population genetics of bryophytes in relation to their reproductive biology. Journal of the Hattori Botanical Laboratory 76: 147–157.
Wyatt, R. & Odrzykoski, I.J. (1998) On the origins of the allopolyploid moss Plagiomnium cuspidatum. The Bryologist 101: 263–271. https://doi.org/10.2307/3244203
Wyatt, R., Odrzykoski, I.J. & Stoneburner, A. (1991) Chemosystematics of the Mniaceae. II. Flavonoids of Plagiomnium Section Rosultata. The Bryologist 94: 443–448. https://doi.org/10.2307/3243842
Wyatt, R., Odrzykoski, I.J. & Stoneburner, A. (1992) Isozyme evidence of reticulate evolution in mosses: Plagiomnium medium is an allopolyploid of P. ellipticum × P. insigne. Systematic Botany 17: 532–550. https://doi.org/10.2307/2419725
Wyatt, R., Odrzykoski, I.J. & Stoneburner, A. (1993a) Isozyme evidence regarding the origins of the allopolyploid moss Plagiomnium curvatulum. Lindbergia 18: 49–58.
Wyatt, R., Odrzykoski, I.J. & Stoneburner, A. (1993b) Isozyme evidence proves that the moss Rhizomnium pseudopunctatum is an allopolyploid of R. gracile × R. magnifolium. Memoirs of the Torrey Botanical Club 25: 21–35.
Wyatt, R., Odrzykoski, I.J. & Stoneburner, A. (2013) Isozyme evidence regarding the nature of polyploidy in the moss genus Cinclidium (Mniaceae). The Bryologist 116: 229–237. https://doi.org/10.1639/0007-2745-116.3.229
Wyatt, R., Odrzykoski, I.J., Stoneburner, A., Bass, H.W. & Galau, G.A. (1988) Allopolyploidy in bryophytes: multiple origins of Plagiomnium medium. Proceedings of the National Academy of Sciences, USA 85: 5601–5604. https://doi.org/10.1073/pnas.85.15.5601