Open Access Open Access  Restricted Access Subscription or Fee Access


A study of molecular sequences, sexuality, and morphological variation in Plagiomnium carolinianum, P. maximoviczii, and P. rhynchophorum (Bryophyta, Mniaceae)



Morphologically, recognition of the genus Plagiomnium may be relatively easy. Yet identifications of closely related species have met great difficulties. The contemporary species delimitations of P. carolinianum, P. maximoviczii, and P. rhynchophorum largely based on sexuality as the sole distinction have not been satisfactory. As shown from literature, character variations among these three taxa were continuous and intergraded within or among different populations throughout a wide geographic range. No gametophytic characters could be reliably used to distinguish them from each other. Molecular phylogenetic analyses using ITS2 and rps4 gene were undertaken to resolve delineations for these three morphologically similar species. The results suggest that they form a well support monophyletic clade, which can be defined as representing one single species with two subspecies, i.e. P. rhynchophorum subsp. maximoviczii and P. rhynchophorum subsp. rhynchophorum. The present molecular study supports the treatment of P. carolinianum as synonym of P. rhynchophorum as purposed previously by Koponen based on morphology.


Asia, China, chromosome number, ITS2, Mniaceae, Mnium, moss, Orthomnion, Plagiomnium, rps4, Bryophytes

Full Text:



Allen, B.H. (2002) Moss flora of Central America, Part 2. Encalyptaceae–Orthotrichaceae. Monographs in Systematic Botany from the Missouri botanical garden 90: 1–699 + viii.

Allen, B.H. (2005) Maine mosses. Sphagnaceae–Timmiaceae. Memoirs of the New York Botanical Garden 93: 1–420 + [xii].

Anderson, L.E. (1954) A new species of Mnium from the southern Appalachians. The Bryologist 57 (3): 177–188.[177:ANSOMF]2.0.CO;2

Bowers, M.C. (1968) A cytotaxonomic study of the genus Mnium in Colorado. Revue bryologique et lichénologique 36: 167-202.

Bowers, M.C. (1980) A cytotaxonomic classification of the Mniaceae (Bryophyta). Lindbergia 6: 22–31.

Brotherus, V.F. (1906[1907]) Orthomniopsis und Okamuraea, zwei neue Laubmoos-gattungen aus Japan. Öfversigt af Finska Vetenskaps-Societetens Förhandlingar 49 (10): 1–4, Tabs. I–II.

Cao, T., Sha, W., Zhang, H.Y. & Xue, H. (2001) Chromosome numbers of 8 species of mosses from northeast China. Guizhou Science 19 (4): 13–15.

Chen, P.C. (1955) Bryophyta nova Sinica. Feddes Repertorium Specierum Novarum Regni Vegetabilis 58: 23–52.

Crum, H. & Anderson, L.E. (1981) Mosses of eastern North America Vol. 1. 663 pp. Columbia University Press, New York.

Eddy, A. (1996) Splachnobryaceae to Leptostomataceae. In: Handbook Malesian Mosses. Vol. 3. Natural History Museum Publications, London, [iv] + 277 pp.

Fedosov, V.E., Fedorova, A.V., Ignatova, E.A., Bobrova, V.K. & Troitsky, A.V. (2015) Rps4 and nad5 sequences indicate the polyphyly of Ditrichaceae and parallelisms in the evolution of haplolepidous mosses. Molecular Biology 49 (6): 890–894.

Gangulee, H.C. & Chatterjee, N.K. (1960) Cytological studies in the mosses of eastern India. II. Nucleus (Calcutta) 3: 165–176.

Grechko, V.V, Fedorova, L.V., Fedorov, A.N., Slobodyanyuk, S.Y., Ryabinin, D.M., Melnikova, M.N., Bannikova, A.A., Lomov, A.A., Sheremeteva, V.A., Gorshkov, V.A., Sevostyanova, G.A. & Semenova, S.K. (1997) Restriction endonuclease analysis of highly repetitive DNA as a phylogenetic tool. Journal of Molecular Evolution 45: 332–336.

Harris, E.S.J. (2008) Paraphyly and multiple causes of phylogenetic incongruence in the moss genus Plagiomnium (Mniaceae). Taxon 57: 417–433.

Hedwig, J. (1801) Species Muscorum Frondosorum. J.A. Barth, Leipzig & Paris, vi + 352 pp.

Hooker, W.J. (1836) Icones Plantarum; or figures with brief descriptive characters and remarks, of new or rare plants, selected from the authors’s herbarium. Vol. 1. London, pl. 1–50 Tabs.

Ignatov, M.S., Afonina, O.M. & Ignatova, E.A. (2006) Checklist of mosses of East Europe and North Asia. Arctoa 15: 1–130.

Inoue, S. & Morita, Y. (1976) Chromosome studies in mosses, VI. Miscellanea Bryologica et Lichénologyca 7: 65–67.

Iwatsuki, Z. & Sharp, A.J. (1968) Some interesting mosses from the Philippines. Journal of the Hattori Botanical Laboratory 31: 50–54.

Kabiersch, W. (1936) Studien über die ostasiatischen arten einiger Laubmoosfamilien (Mniaceae-Bartramiaceae). Hedwigia 76: 1–94.

Kindberg, N.C. (1897) Part 2. Acrocarpous. In: European and North American Bryineae Linköpings Lithografiska. Linköping, pp. 153–410.

Koponen, T.J. (1968) Generic revision of Mniaceae Mitt. (Bryophyta). Annales Botanici Fennici 5: 117–151.

Koponen, T. (1971) A preliminary report on Plagiomnium sect. Rostrata in Asia. Hikobia 6: 57–58.

Koponen, T.J. (1972) The East Asiatic species of Plagiomnium sect. Rostrata (Bryophyta). Acta Botanica Fennica 97: 3–29.

Koponen, T. (1977) Miscellaneous notes on Mniaceae (Bryophyta). I. New combinations. Annales Botanici Fennici 14: 6.

Koponen, T.J. (1980) A synopsis of Mniaceae (Bryophyta). II. Orthomnion. Annales Botanici Fennici 17: 35–55.

Koponen, T.J. (1981a) A synopsis of Mniaceae (Bryophyta). V. Taxa in Africa south of the Sahara. Annales Botanici Fennici 18: 105–111.

Koponen, T.J. (1981b) Miscellaneous notes on Mniaceae (Bryophyta). VIII. Verification of the vouchers of some chromosome numbers reports. Annales Botanici Fennici 18: 73–82.

Koponen, T. (1981c) A synopsis of Mniaceae (Bryophyta). VI. Southeast Asian taxa. Acta Botanica Fennica 117: 1–34.

Koponen, T.J. (2014) Bryophyte flora of Hunan Province, China. 18. Mniaceae subfam. Mnioideae (Musci). Acta Bryolichenologica Asiatica 5: 39–72.

Koponen, T.J. & Sun, Y. (2017[2016]) Preliminary study on phylogenetic position and delimitation of the ciliate arthrodontous genera of the moss family Mniaceae. Journal of Bryology 39 (1): 23–38.

Larkin, M.A., Blackshields, G, Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F.C., Wallace, I.M., Wilm, A., Lopez, R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23 (21): 2947–2948.

Li, X.J. (2006) Mniaceae. In: Li, X.J. (Ed.) Flora Bryophytorum Sinicorum. Vol. 4. Science Press, Beijing, pp. 111–178.

Li, X.J., He, S. & Zang, M. (2007) Mniaceae. In: Li, X.J., Crosby, M.R. & He, S. (Eds.) Moss Flora of China, English Version. Vol. 4. Science Press & Missouri Botanical Garden, Beijing, New York & St. Louis., pp. 93–144.

Lindberg, S.O. (1872 [1873]) Contributio ad Floram Cryptogamam Asiae Boreali-Orientalis. Societatis Litterariae Fennicae, Helsingfors, pp. 221–280.

Lowry, R.J. (1948) A cytotaxonomic study of the genus Mnium. Memoirs of Torrey Botanical Club 20: 1–42.

McIntosh, T.T. (2014) Mniaceae, pp. 215–243. In Flora of North America editorial committee (Eds.) Bryophyta, part 2. Flora of North America 28: i–xxii, 1–702.

Mitten, W. (1859) Musci Indiae Orientalis, an enumeration of the mosses of the East Indies. Journal of the Proceeding of the Linnean Society, Botany 1: 1–96.

Noguchi, A. (1959) Mosses from Pakistan, 2. Journal of the Hattori Botanical Laboratory 21: 292–295.

Noguchi, A. (1960) Mnium cuspidatum and M. trichomanes in Japan. Miscellanea Bryologica et Lichenologica 2: 1–2.

Noguchi, A. (1966) Musci. In: Flora of Eastern Himalaya. University Museum, University of Tokyo, Tokyo, pp. 537–591.

Noguchi, A. (1973) Mosses of Ceylon collected by Dr. Hiroshi Inoue. Bulletin of the National Science Museum, Tokyo New series, 16: 305–316.

Noguchi, A. (1989) Illustrated Moss Flora of Japan. Part 3. Hattori Botanical Laboratory, Nichinan, pp. 492–742.

Ochyra, R., Bednarek-Ochyra, H.K. & Plášek, V. (2017) Proposal to conserve the name Plagiomnium against Orthomnion and Orthomniopsis (Bryophyta: Mniaceae). Taxon 66 (5): 1218–1220.

Oguri, E., Yamaguchi, T., Tsubota, H., Shimamura, M. & Deguchi, H. (2006) Morphological and molecular analyses to solve a taxonomical controversy of Leucobryum juniperoideum (Brid.) Müll. Hal. and L. humillimum Cardot (Leucobryaceae, Musci) in Japan. Hikobia 14: 387–398.

Oguri, E., Yamaguchi, T., Tsubota, H., Shimamura, M. & Deguchi, H. (2010) Phylogenetic relationships among Leucobryum juniperoideum and related species (Leucobryaceae, Musci) inferred from ITS sequences. Hikobia 15 (4): 453–462.

Ono, K. (1970) Karyological studies on Mniaceae and Polytrichaceae, with special reference to the structural sex-chromosomes 2. Journal of Science of the Hiroshima University Series B, Division 2 (Botany), 13: 107–166.

Ono, K., Morita, Y., Taguchi, M. & Inoue, S. (1977) On the chromosome number in Mniaceae. Kumamoto Journal of Science Section 2, Biology, 13: 49–65.

O’Shea, B.J. (2002) Checklist of the mosses of Sri Lanka. Journal of the Hattori Botanical Laboratory 92: 125–164.

Palisot de Beauvois, A.M.F.J. (1805) Prodrome des Cinquième et Sixième Familles de I’Aethéogam. Pournier Fils, Paris, ii + 114 pp.

Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 (12): 1572–1574.

Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28 (10): 2731–2739.

Tatuno, S. & Ono, K. (1966) Zytologische Untersuchungen über die Arten von Mniaceae aus Japan. Journal Hattori Botanical Laboratory 29: 79–95.

Tian, X.H., Xiao, Y.P., Liu, Q.H., Zhu, B.C., Wang, Z.H. & Yuan, S.Q. (1994) Observations on chromosomes of fourteen moss species from the Qinling Range, China. Acta Phytotaxonomica Sinica 32 (3): 240–245.

Tixier, P. (1971) Bryophytae Indosinicae. Mousses de Thailande. Annales de la Faculté des Sciences, Université de Phnom Penh 4: 91–166.

Tixier, P. (1975) A contribution to the knowledge of the mountain moss flora of Sri Lanka. Ceylon Journal of Science 11: 123–134.

Tixier, P. & Smitinand, T. (1966) Checklist of the moss collection in the Forest Herbarium, Royal Forest Department, Bangkok. Natural History Bulletin of the Siam Society 21 (3–4): 161–195.

Uniyal, P.L. (1997) Cytological studies on Plagiomnium T. Kop. of western Himalaya. Acta Botanica Indica 25: 37–42.

Vanderpoorten, A., Boles, S. & Shaw, A.J. (2003) Patterns of molecular and morphological variation in Leucobryum albidum, L. glaucum, and L. juniperoideum (Bryopsida). Systematic Botany 28: 651–656.

Wilson, W. (1857) Enumeration of the mosses collected in India by Dr. J. D. Hooker, F.R.S., and Dr. Thomas Thomson, F.R.S., with their habitats, elevations, and the numbers under which they have been distributed; arranged by W. Mitten, Esq. Hooker’s Journal Botany Kew Garden Miscellany 9: 363–370.

Withering, W. (1801) A Systematic Arrangement of British Plants (ed. 4) 3. pp. i–iii, 445–900.

Wynns, J.T. & Lange, C.B.A. (2014) A comparison of 16 DNA regions for use as phylogenetic markers in the pleurocarpous moss genus Plagiothecium (Hypnales). American Journal of Botany 101 (4): 652–669.


  • There are currently no refbacks.

ISSN 1179-3155 (print); ISSN 1179-3163 (online)

Published by Magnolia Press, Auckland, New Zealand