Open Access Open Access  Restricted Access Subscription or Fee Access


Towards a monophyletic classification of Lejeuneaceae I: subtribe Leptolejeuneinae subtr. nov.



We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region of seven accessions of Leptolejeunea to explore its phylogenetic position. Maximum parsimony and maximum likelihood analyses led to similar topologies but deeper nodes received good bootstrap support only with maximum likelihood methods. Leptolejeunea formed an early diverging, robust monophyletic lineage within Lejeuneaceae tribe Lejeuneeae. Contrary to earlier belief, it is not closely related to Drepanolejeunea. To amend the current classification of Lejeuneeae into subtribes, we propose Leptolejeuneinae, subtr. nov.


epiphyll, Lejeuneeae, Leptolejeunea, liverwort, Porellales, taxonomy

Full Text:

PDF/A (385.57 KB)


Bischler, H. (1969) Le genre Leptolejeunea (Spruce) Steph. en Amérique. Nova Hedwigia 17: 255-350.

Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModeltest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772.

Dong, S., Schäfer-Verwimp, A., Meinecke, P., Feldberg, K., Bombosch, A., Pócs, T., Schmidt, A.R., Reitner, J., Schneider, H. & Heinrichs, J. (2012) Tramps, narrow endemics and morphologically cryptic species in the epiphyllous liverwort Diplasiolejeunea. Molecular Phylogenetics and Evolution 65: 582–594.

Dong, S., Schäfer-Verwimp, A., Pócs, T., Feldberg, K., Czumaj, A., Schmidt, A.R., Schneider., H. & Heinrichs, J. (2013) Size doesn´t matter – recircumscription of Microlejeunea based on molecular and morphological evidence. Phytotaxa 85: 41–55.

Dumortier, B.C.J. (1831) Sylloge Jungermannidearum Europae indigenarum: earum genera et species systematice complectens. Casterman, Tournay, 100 pp.

Dumortier, B.C.J. (1835) Recueil d'Observations sur les Jungermanniacées. Blanquart, Tournay, 27 pp.

Gradstein, S.R. (2013) A classification of Lejeuneaceae based on molecular and morphological evidence. Phytotaxa 100: 6–20.

Gradstein, S.R., Churchill, S.P. & Salazar-Allen, N. (2001) Guide to the bryophytes of tropical America. Memoirs of the New York Botanical Garden 86: 1–577.

Gradstein, S.R., Reiner-Drehwald, M.E. & Schneider, H. (2003) A phylogenetic analysis of the genera of Lejeuneaceae (Hepaticae). Botanical Journal of the Linnean Society 143: 391–410.

Gradstein, S.R., Wilson, R., Ilkiu-Borges, A.L. & Heinrichs, J. (2006) Phylogenetic relationships and neotenic evolution of Metzgeriopsis (Lejeuneaceae) based on chloroplast DNA sequences and morphology. Botanical Journal of the Linnean Society 151: 293–308.

Grolle, R. (1976) Drepanolejeunea subgen. Kolpolejeunea – eine neue Untergattung aus der Paläotropis. Journal of the Hattori Botanical Laboratory 40: 191–216.

Grolle, R. (1980) Schusterolejeunea Grolle nom. nov. statt Cladocolea Schust. 1963, non Van Tieghem 1895. Journal of Bryology 11: 105–106.

Grolle, R. (1995) The Hepaticae and Anthocerotae of the East African Islands. Bryophythorum Bibliotheca 48: 1–178.

Guindon, S. & Gascuel, O. (2003) A simple, fast and accurate method to estimate large phylogenies by maximum likelihood. Systematic Biology 52: 696–704.

Hall, T.A. (1999) BIOEDIT: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposia Series 41: 95–98.

Hartmann, F.A., Wilson, R., Gradstein, S.R., Schneider, H. & Heinrichs, J. (2006) Testing hypotheses on species delimitations and disjunctions in the liverwort Bryopteris (Jungermanniopsida: Lejeuneaceae). International Journal of Plant Sciences 167: 1205–1214.

Hattori, S. (1951) Contributio ad Floram Hepaticarum Yakusimensem, V. Journal of the Hattori Laboratory 5: 43–68.

He, Q. & Zhu, R.-L. (2011) Spore output in selected species of Lejeuneaceae. Cryptogamie, Bryologie 32: 107–112.

Heinrichs, J., Czumaj, A., Dong, S., Scheben, A., Schäfer-Verwimp, A., Peralta, D.F., Feldberg, K., Schmidt, A.R. & Schneider, H. (2014) The Bromeliaceae tank dweller Bromeliophila (Lejeuneaceae, Porellales) is a member of the Cyclolejeunea-Prionolejeunea clade. Plant Systematics and Evolution 300: 63–73.

Heinrichs, J., Dong, S., Feldberg, K., Schäfer-Verwimp, A. & Schmidt, A.R. (2012a) Sphaerolejeunea (Lejeuneaceae, Porellales) is a synonym of Lejeunea. Phytotaxa 69: 7–15.

Heinrichs, J., Dong, S., Schäfer-Verwimp, A., Pócs, T., Feldberg, K., Czumaj, A., Schmidt, A.R., Reitner, J., Renner, M.A.M., Hentschel, J., Stech, M. & Schneider, H. (2013) Molecular phylogeny of the leafy liverwort Lejeunea (Porellales): Evidence for a Neotropical origin, uneven distribution of sexual systems and insufficient taxonomy. PloS ONE 8(12): e82547.

Heinrichs, J., Dong, S., Yu, Y., Schäfer-Verwimp, A., Pócs, T., Feldberg, K., Hentschel, J., Schmidt, A.R. & Schneider, H. (2012b) A 150 year old mystery solved: Transfer of the rheophytic liverwort Myriocolea irrorata to Colura. Phytotaxa 66: 55–64.

Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing the confidence in phylogenetic analysis. Systematic Biology 42: 182–192.

Mason-Gamer, R.J. & Kellogg, E.A. (1996) Testing for phylogenetic conflict among molecular data sets in the tribe Triticeae (Gramineae). Systematic Biology 45: 524–545.

Mitten, W. (1861) Hepaticae Indiae Orientalis: an ennumeration of the Hepaticae of the East Indies. Journal of the Proceedings of the Linnean Society, Botany 5: 89–128.

Pócs, T. (1996) Epiphyllous liverwort diversity at worldwide level and its threat and conservation. Anales del Instituto de Biológia de la Universidad Nacional Autónoma de México, Serie Botánica 67: 109–127.

Schiffner, V. (1893) Hepaticae. In: Engler, A. & Prantl, K., Die natürlichen Pflanzenfamilien 1. Engelmann, Leipzig, pp. 97–141.

Schiffner, V. (1898) Conspectus Hepaticarum Archipelagi Indici. Staatsdruckerei, Batavia, 425 pp.

Schuster, R.M. (1963) An annotated synopsis of the genera and subgenera of Lejeuneaceae. Beihefte zur Nova Hedwigia 9: 1–203.

Schuster, R.M. & Giancotti, C. (1993) On Vitalianthus Schust. & Giancotti, a new genus of Lejeuneaceae. Nova Hedwigia 57: 445–456.

Spruce, R.M. (1884) Hepaticae Amazonicae et Andinae. Tribus I: Jubuleae. Transactions and Proceedings of the Botanical Society of Edinburgh 15: 1–308.

Swofford, D.L. (2000) PAUP*, phylogenetic analyses using parsimony (* and other methods), version 4.01b10. Sinauer Associates, Sunderland, Massachusetts.

Wilson, R., Gradstein, S.R., Heinrichs, J., Groth, H., Ilkiu-Borges, A.L. & Hartmann, F.A. (2004) Phylogeny of Lejeuneaceae: A cladistic analysis of chloroplast gene rbcL and morphology with preliminary comments on the mitochondrial nad4-2 spacer region. Monographs in Systematic Botany from the Missouri Botanical Garden 98: 189–202.

Wilson, R., Gradstein, S.R., Schneider, H. & Heinrichs, J. (2007a) Unravelling the phylogeny of Lejeuneaceae (Jungermanniopsida): evidence for four main lineages. Molecular Phylogenetics and Evolution 43: 270–282.

Wilson, R., Heinrichs, J., Hentschel, J., Gradstein, S.R. & Schneider, H. (2007b) Steady diversification of derived liverworts under Tertiary climatic fluctuations. Biology Letters 3: 566–569.

Ye, W., Wei, Y.M., Schäfer-Verwimp, A. & Zhu, R.L. (2013) Phylogenetic position of Oryzolejeunea (Lejeuneaceae, Marchantiophyta): Evidence from molecular markers and morphology. Journal of Systematics and Evolution 51: 468–475.

Yu, Y., Pócs, T., Schäfer-Verwimp, A., Heinrichs, J., Zhu, R.-L. & Schneider, H. (2013) Evidence for rampant homoplasy in the phylogeny of the epiphyllous liverwort genus Cololejeunea (Lejeuneaceae). Systematic Botany 38: 553–564.

Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. GARLI version 2.0 available online at


  • There are currently no refbacks.

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

Published by Magnolia Press, Auckland, New Zealand