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L.D. Alcaraz et al. (2018): Marchantia liverworts as a proxy to plants’ basal microbiomes. Open access, Scientific Reports, 8.

! Australian National Botanic Gardens and Centre for Australian National Biodiversity Research, Canberra: Bryophytes - the world of hornworts, liverworts and mosses.

R.S. Barclay et al. (2013): New methods reveal oldest known fossil epiphyllous moss: Bryiidites utahensis gen. et sp. nov.(Bryidae). In PDF, American Journal of Botany, 100: 2450-2457.
Using the zstacking software package Automontage (Syncroscopy, Cambridge, UK).

Michael Becker: Moose (in German).

Museum of Paleontology (UCMP), University of California, Berkeley: Introduction to the Bryophyta.

A.C. Bippus (2018): Extending the fossil record of the Polytrichaceae (Bryophyta): insights from the early Cretaceous of Vancouver Island, Canada. In PDF, thesis, Humboldt State University, Arcata, Canada. See also here.

A.C. Bippus et al. (2017): Extending the fossil record of Polytrichaceae: Early Cretaceous Meantoinea alophosioides gen. et sp. nov., permineralized gametophytes with gemma cups from Vancouver Island. In PDF, American Journal of Botany, 104: 584–597. See also here.

M.I. Bidartondo et al. (2011): The dawn of symbiosis between plants and fungi. In PDF, Biology Letters. See also here.

! Bildatlas der Moose Deutschlands (by Michael Lueth, Freiburg, Germany). Habitat and bryophyte photographs. Excellent! This is a comprehensive discussion publishing site specially developed for school and college students, teachers, and learners.
Go to:
List of 4 Important Fossil Brophytes.

B. Bomfleur et al. (2014): Diverse bryophyte mesofossils from the Triassic of Antarctica. In PDF, see also here.

J.L. Bowman et al. (2017): Insights into Land Plant Evolution Garnered from the Marchantia polymorpha Genome. Abstract, Cell, 171: 287-304. See also here (in PDF).

Jamie Boyer, The New York Botanical Garden: The Paleoplant Website. An educational resource for students and teachers studying Earth's history, fossils, and evolution.
! Go to: Bryophytes.

The British Bryological Society:
What is a bryophyte?
Bryological Journals. A link directory to journals devoted specifically to bryophytes (sometimes also including lichens) (34 journals), journals with a regular bryological content (16 journals), and to 'Extinct' bryological journals (31 journals).
World Wide Bryology. Links to other sites.

W.R. Buck et al. (2012): Recent literature on bryophytes—115(3). The Bryologist, 115.

! Southern Illinois University, Carbondale: Bryophytes.

C.M. Sean Carrington: The Bryophytes.
Snapshot provided by the Internet Archive´s Wayback Machine.

! Michael Clayton, Department of Botany, University of Wisconsin, Madison: Plant Teaching Collection. Some image collections. Excellent! Go to:

C. Cardona-Correa et al. (2016): Peat Moss–Like Vegetative Remains from Ordovician Carbonates. Free access, International Journal of Plant Sciences, 177: 523-538.

! B.J. Crandall-Stotler and S.E. Bartholomew-Began (2007): Morphology of Mosses (Phylum Bryophyta). In PDF. Excellent!
In: Flora of North America Editorial Committee (eds): Flora of North America North of Mexico. Volume 27: Bryopsida: Mosses, Part 1, 3–13 pp. Oxford University Press, New York.
This expired link is available through the Internet Archive´s Wayback Machine.

C. DeLavoi: Bryology. Beautiful photographs.

C.F. Delwiche et al. (2017): Land Plant Model Systems Branch Out. In PDF, Cell, 171.
"Liverworts may be the sister taxon to all other land plants, and the genome shows features that illuminate the ancestor of all land plants and give insights into how plant systems function and evolved".

Deutsche Apotheker Zeitung (in German): Moose – vielversprechend und enttäuschend.

D. Edwards (2000): The role of Mid-Palaeozoic mesofossils in the detection of early bryophytes. In PDF, Philosophical Transactions of the Royal Society B.

K. Feldberg et al. (2017): Problems related to the taxonomic placement of incompletely preserved amber fossils: transfer of the Paleogene liverwort Cylindrocolea dimorpha (Cephaloziellaceae) to the extant Odontoschisma sect. Iwatsukia (Cephaloziaceae). In PDF, Foss. Rec., 20: 147–157.

K. Feldberg et al. (2013): Exploring the impact of fossil constraints on the divergence time estimates of derived liverworts. In PDF, Plant Syst. Evol., 299: 585-601. See also here.

B.J. Fletcher et al. (2008): Atmospheric carbon dioxide linked with Mesozoic and early Cenozoic climate change. In PDF, Nat. Geosci., 1: 43-48.

B.J. Fletcher et al. (2005): Fossil bryophytes as recorders of ancient CO2 levels: Experimental evidence and a Cretaceous case study. In PDF.

J.R. Flores et al. (2017): Combined phylogenetic analysis of the subclass Marchantiidae (Marchantiophyta): towards a robustly diagnosed classification. Abstract, Cladistics, 1–25. See also here or there (in PDF).

L.L. Forrest et al. (2006): Unraveling the evolutionary history of the liverworts (Marchantiophyta): multiple taxa, genomes and analyses. Abstract, The Bryologist, 109: 303-334.

Fossil Hunters (by Dennis Larson):
Fossil Plants.
Marchantiophytina (September 06, 2018).

! J.M. Glime (2017): Adaptive Strategies: Spore Dispersal Vectors. Chapt. 4-9. In: Glime, J. M. Bryophyte Ecology. Volume 1. 4-9-1 Physiological Ecology. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 6 March 2017. See also here.

! J.M. Glime (2013): Meet the Bryophytes. In PDF, Chapt. 2-1. In: Glime, J. M. Bryophyte Ecology. Volume 1. Physiological Ecology. Ebook 2-1-1 sponsored by Michigan Technological University and the International Association of Bryologists.

! J.M. Glime (2007): Bryophyte Ecology. Ebook sponsored by Department of Biological Sciences, Michigan Technological University and the International Association of Bryologists. Excellent!

! L.E. Graham et al. (2010): Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats. In PDF, American Journal of Botany, 97: 268-275. See also:
! T.N. Taylor et al. (2010): The enigmatic Devonian fossil Prototaxites is not a rolled-up liverwort mat: Comment on the paper by Graham et al.(AJB 97: 268-275). In PDF. See also:
! L.E. Graham et al. (2010): Rolled liverwort mats explain major Prototaxites features: Response to commentaries.

L.E. Graham et al. (2004): Resistant tissues of modern marchantioid liverworts resemble enigmatic Early Paleozoic microfossils. In PDF, PNAS, 101: 11025-11029.

Tomas Hallingbäck and Nick Hodgetts (2000), IUCN/SSC Bryophyte Specialist Group (International Union for Conservation of Nature):
! Mosses, Liverworts and Hornworts. In PDF, 11 MB. As well as the threats and importance of bryophytes, this document explains their habitats and classification details.
Slow download. Snapshots provided by the Internet Archive´s Wayback Machine.

Botanik Online, Hamburg: Bryophyta (Moose) (in German). Snapshot taken by the Internet Archive´s Wayback Machine.

! J. Heinrichs et al. (2015): Molecular and Morphological Evidence Challenges the Records of the Extant Liverwort Ptilidium pulcherrimum in Eocene Baltic Amber. Open access, PLoS ONE 10: e0140977.

J. Heinrichs et al. (2014): A Jurassic moss from Northeast China with preserved sporophytes. In PDF, see also here.

J. Heinrichs et al. (2012): The sporophyte of the Paleogene liverwort Frullania varians Caspary. In PDF, Fossil Record, 15: 115–120.

J. Heinrichs et al. (2011): Kaolakia borealis nov. gen. et sp. (Porellales, Jungermanniopsida): A leafy liverwort from the Cretaceous of Alaska. In PDF, Review of Palaeobotany and Palynology.

! J. Heinrichs et al. (2007): Evolution of leafy liverworts (Jungermanniidae, Marchantiophyta): estimating divergence times from chloroplast DNA sequences using penalized likelihood with integrated fossil evidence. Abstract, Taxon, 56: 31–44. See also here (in PDF).

! L.V. Hernick et al. (2008): Earth's oldest liverworts — Metzgeriothallus sharonae sp. nov. from the Middle Devonian (Givetian) of eastern New York, USA. Abstract, Review of Palaeobotany and Palynology, 148.

! Rolf Holderegger and Ariel Bergamini, Eidgenössische Forschungsanstalt WSL, Biodiversität & Naturschutzbiologie (ETH Zürich):
Biologie der Moose und Farne, 1. Moose.
Lecture notes (in German).

! M. Hübers and H. Kerp (2012): Oldest known mosses discovered in Mississippian (late Visean) strata of Germany. In PDF, Geology.

C. Humphreys: Fossil bryophyte proxy contributes to palaeo-atmospheric CO2 predictions. In PDF. See also here.

C. Humphreys: Literature Review: Abiotic and Biotic Influences on the Productivity of Early Land Plants. In PDF. See also here.

M. S. Ignatov et al. (2012): New Bryokhutuliinia species (Bryophyta) with sporophytes from the Upper Jurassic of Transbaikalia. In PDF, Arctoa, 21: 69-78.

J.A. Janssens (2014): Field Guide to Mosses and Liverworts of Minnesota's Calcareous Fens. In PDF.

Joannes A. Janssens, Minnesota Ground Water Association: Introduction to Bryology. Powerpoint presentation. See also here.
Snapshot provided by the Internet Archive´s Wayback Machine.

J.A. Janssens (1990): Methods in Quaternary Ecology 11. Bryophytes. In PDF, Geoscience Canada, 17.

T. Katagiri and A. Hagborg (2015): Validation of ordinal and family names for a Triassic fossil liverwort, Naiadita. (Naiaditaceae, Marchantiopsida). In PDF, Phytotaxa, 222: 165-166.

! K.-P. Kelber (2019): Naiadita lanceolata (Marchantiophyta) from the Middle Triassic (Ladinian) of Germany: a new reconstruction attempt and considerations on taphonomy. Abstract, PalZ, 93: 499-515.

Michael Knee, Ohio State University: The Bryophyta. A version archived by Internet Archive Wayback Machine.

A.S Konopka, P.S. Herendeen and P.R. Crane (1998): Sporophytes and gametophytes of Dicranaceae from the Santonian (Late Cretaceous) of Georgia, USA. American Journal of Botany, 85.
This expired link is now available through the Internet Archive´s Wayback Machine.

Ari Kornfeld, Natural Perspective: The Plant Kingdom, Mosses and Allies. Snapshot taken by the Internet Archive´s Wayback Machine.

J. Kowal et al. (2018): From rhizoids to roots? Experimental evidence of mutualism between liverworts and ascomycete fungi. In PDF, Annals Of Botany, 121: 221-227. See also here.

Valentin A. Krassilov (1987): Palaeobotany of the mesophyticum: state of the art. In PDF, Review of Palaeobotany and Palynology, 50: 231-254. A version archived by Internet Archive Wayback Machine.

Shawn Krosnick and Kevin E. Indoe, The New York Botanical Garden: What is a bryophyte anyway?

C.C. Labandeira et al. (2014): Middle Devonian liverwort herbivory and antiherbivore defence. In PDF, New Phytologist, 202: 247–258. See also here.

W.S. Lacey (1969): Fossil bryophytes. Abstract, Biological Reviews.

B. Laenen et al. (2016): Geographical range in liverworts: does sex really matter? In PDF, Journal of Biogeography, 43: 627–635. See also here (abstract).

! B. Laenen et al. (2014): Extant diversity of bryophytes emerged from successive post-Mesozoic diversification bursts. In PDF. See also here (abstract).

R. Li et al. (2019): Ricciopsis sandaolingensis sp. nov., a new fossil bryophyte from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang, Northwest China. In PDF, Palaeontologia Electronica, 22.2.42A 1-11. See also here.

R. Li et al. (2016): Fossil Liverworts from the Lower Cretaceous Huolinhe Formation in Inner Mongolia, China. In PDF, slow download. See also here (abstract).

R. Li et al. (2014): Marchantites huolinhensis sp. nov. (Marchantiales) - A new fossil liverwort with gemma cups from the Lower Cretaceous of Inner Mongolia, China. In PDF, Cretaceous Research, 50: 16-26.

! R. Ligrone et al. (2012): Major transitions in the evolution of early land plants: a bryological perspective. In PDF, Annals of botany.

Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima.
This expired link is now available through the Internet Archive´s Wayback Machine.

X. Liu et al. (2018): Liverwort Mimesis in a Cretaceous Lacewing Larva. Open access, Current Biology, 28: 1475-1481.
Note figure 3: Reconstruction of two larvae resting on liverworts.

Y.S. Mamontov and M.S. Ignatov (2019): How to rely on the unreliable: Examples from Mesozoic bryophytes of Transbaikalia. In PDF, Journal of Systematics and Evolution. See also here.

E.V. Maslova et al. (2012): On the leaf development in Palaeozoic mosses of the order Protosphagnales. In PDF, Arctoa, 21: 241-264.

D. Mauquoy et al. (2010): A protocol for plant macrofossil analysis of peat deposits. PDF file, Mires and Peat, 7.

S. McLoughlin and C. Strullu-Derrien (2015): Biota and palaeoenvironment of a high middle-latitude Late Triassic peat-forming ecosystem from Hopen, Svalbard archipelago. In PDF. Permineralized bryophyte sporangium!

! H.A. Miller (1982): Bryophyte evolution and geography. In PDF, Biological Journal of the Linnean Society.

Missouri Botanical Garden, St. Louis:
! Index of Mosses Database (W3MOST). This database of moss names was prepared as part of the Index of Mosses project at the Missouri Botanical Garden, providing guides to newly published nomenclature for mosses. See also:
Bryology Research. Annotated links.

! Missouri Botanical Garden, St. Louis: Bryological Glossary. GLOSSARIUM POLYGLOTTUM BRYOLOGIAE: A multilingual glossary for bryology. Originally published in Monographs in Systematic Botany from the Missouri Botanical Garden Vol. 33, 297 pp. 1990. The multilingual Glossary was created so that bryologists and others could avoid misinterpretations of literature, and general confusion resulting from a lack of standardized terms. The terms are listed alphabetically, and include Latin entries, as well as French, German, and Spanish translations. Excellent!

! J.L. Morris et al. (2018): The timescale of early land plant evolution. In PDF, PNAS, 115. See also here.

! Terence M. Murphy, Thomas L. Rost and Michael G. Barbour (2015), University of California, Davis, CA:
Plant Biology. Lecture notes, in PDF. Please take notice: Book announcement.
See for instance: Bryophytes.
The Early Tracheophytes.

New World Encyclopedia:

The New York Botanical Garden: American Bryophyte Catalog.

D.L. Nickrent et al. (2000): Multigene phylogeny of land plants with special reference to bryophytes and the earliest land plants. PDF file, Molecular Biology and Evolution: 17: 1885-1895. A version archived by Internet Archive Wayback Machine.

! Dan Nickrent, Kevin C. Nixon & Dale Vitt (Curatorial Board; website served from Southern Illinois University, Carbondale): Phyto Images. This site includes a wide variety of vascular plant and bryophyte photos of high quality. Phyto Images belongs to DOL (, which is a web interface based on the Encino Software Project. The Encino project is a unified set of software tools for storing, retrieving, and analyzing biodiversity. Search the Cladogramm Database or use the Diagnostic Keys. Superbly done!

Karl J. Niklas & Tom Silva, Department of Plant Biology, Cornell University, Ithaca, NY: Introductory Botany. Review Topics, Review of Algae, Bryophytes, Pteridophytes- Common Links Between Each Group of Plants.
These expired links are available through the Internet Archive´s Wayback Machine.

Silvia Pressel et al. (2010): Fungal symbioses in bryophytes: New insights in the Twenty First Century. PDF file, Phytotaxa, 9: 238-253. See also here (open access).

! Y.-L. Qiu et al. (2006): The deepest divergences in land plants inferred from phylogenomic evidence. In PDF, PNAS, 103: 15511-15516

J. Quirk et al. (2015): Constraining the role of early land plants in Palaeozoic weathering and global cooling. Proc. R. Soc., B 282.

! K.S. Renzaglia et al. (2018): Morphology supports the setaphyte hypothesis: mosses plus liverworts form a natural group. Abstract, See also here (in PDF).

K.S. Renzaglia et al. (2017): Hornwort stomata: architecture and fate shared with 400 million year old fossil plants without leaves. In PDF, Plant Physiology, 177. See also here.

W.R. Rimington et al. (2018): Ancient plants with ancient fungi: liverworts associate with early-diverging arbuscular mycorrhizal fungi. Proc. R. Soc. B, 285: 20181600. See also here.

Department of Life Science, Riverside City College, Riverside, CA: Botany lecture notes. Powerpoint presentations. e.g. to:
Introduction to the Plant Kingdom: Bryophytes.

J.P. Rose et al. (2016): Shape analysis of moss (Bryophyta) sporophytes: Insights into land plant evolution. Am. J. Bot., 103: 652-662. See also here.

! Malcolm L. Sargent, Department of Plant Biology, University of Illinois, Urbana, IL, and Diane H. Lucas, Wellington, OH:
A Perspective Oriented Guide for the Identification of North American Bryophyte Genera. The goal of this guide is to give perspective on the characters used in the identification of mosses.
! Don´t miss the useful Glossary. See also here.

A. Savoretti et al. (2018): Grimmiaceae in the Early Cretaceous: Tricarinella crassiphylla gen. et sp. nov. and the value of anatomically preserved bryophytes. Free access, Annals of Botany, 121: 1275–1286. See also here.
"... One fossil moss gametophyte preserved in a carbonate concretion was studied in serial sections prepared using the cellulose acetate peel technique. ..."

H. Schneider (2019): Celebrating Research Devoted to Seed-Free Land Plants. Free access, Journal of Systematics and Evolution, 57: 303-304. See also here (table of contents, free access).

K.W. Schwab et al. (2011): Dwarf Hepatic Floral Fragments (Possibly Liverworts) from the Upper Mississippian Barnett Shale in the Fort Worth Basin of North Central Texas. In PDF.

! A.J. Shaw et al.(2011): Bryophyte diversity and evolution: Windows into the early evolution of land plants. In PDF, Am. J. Bot., 98: 352-369. See also here (abstract).

! J. Shaw and K. Renzaglia (2004): Phylogeny and diversification of Bryophytes. Free access, Amer. J. Botany, 91: 1557-1581. See also here.

G.W.K. Shelton et al. (2016): Krassiloviella limbelloides gen. et sp. nov.: Additional Diversity in the Hypnanaean Moss Family Tricostaceae (Valanginian, Vancouver Island, British Columbia). In PDF, Int. J. Plant Sci., 177: 792–808. See also here (abstract).
Fossils are preserved anatomically in carbonate concretions and studied in serial sections prepared using the cellulose acetate peel technique.

G.W.K. Shelton et al. (2015): Exploring the fossil history of pleurocarpous mosses: Tricostaceae fam. nov. from the Cretaceous of Vancouver Island, Canada. In PDF, American Journal of Botany. See also here. Lexikon der Biologie, Moose (in German).

C.M. Steenbock et al. (2011): A new family of leafy liverworts from the middle Eocene of Vancouver Island, British Columbia, Canada. Abstract, American Journal of Botany, 98.

! Raymond E. Stotler and Barbara J. Crandall-Stotler, Department of Plant Biology, Southern Illinois University, Carbondale: Bryophytes. This website is a resource devoted to Bryology, the branch of plant science concerned with the study of mosses, liverworts and hornworts. It provides information on the classification, structural features, natural history, ecology and evolutionary relationships of these plants.

Y. Sun (2019): Phylogeny and biogeography of liverworts Marchantiophyta evidence from the Southern Hemisphere family Schistochilaceae and the cosmopolitan genus Herbertus Herbertaceae. In PDF, Thesis, University of Helsinki, Faculty of Biological and Environmental Sciences Doctoral Program in Plant Sciences, Finnish Museum of Natural History, Botany Unit. See also here.

! E.L. Taylor and T.N. Taylor (2012): Paleozoic mosses: Small, but no longer inconspicuous. In PDF, Geology, 40: 767-768.

! A.M.F. Tomescu et al. (2018): Why Are Bryophytes So Rare in the Fossil Record? A Spotlight on Taphonomy and Fossil Preservation Transformative Paleobotany. Abstract. In: Papers to Commemorate the Life and Legacy of Thomas N. Taylor. Pages 375-416. See also here (in PDF).
! A.M.F. Tomescu et al. (2017): The bryophyte fossil record database, Paleozoic through Paleogene. Zip-file (doc), hosted by Book companion - Transformative Paleobotany.
"The tables contain mosses and liverworts and hornworts, respectively, arranged in alphabetical order. Each entry represents a taxonomically and stratigraphically distinct (i.e., in terms of rock unit) occurrence".

A.M.F.M. Tomescu (2018): Exquisitely preserved tiny fossils are key for understanding moss evolution. Botany One.

A.M.F. Tomescu (2016): The Early Cretaceous Apple Bay flora of Vancouver Island: a hotspot of fossil bryophyte diversity. In PDF, Botany, 9. See also here.

! Alain Vanderpoorten and Bernard Goffinet (2009): Introduction to Bryophytes, Evolutionary significance of bryophytes. In PDF, Cambridge University Press. See also here.

A. Villarreal et al. (2015): Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate. Abstract, New Phytologist. See also here (in PDF).

Morgan Vis and Emily Miller, Ohio University: Bryophyte Home Page.

! Matt von Konrat et al. (2010): A special issue of Phytotaxa dedicated to Bryophytes: The closest living relatives of early land plants. Editorial (PDF), Phytotaxa, 9: 5-10. Go to: Table of Contents (open access). See especially:
Matt von Konrat et al. (2010): Early Land Plants Today (ELPT): How many liverwort species are there? PDF file, Phytotaxa, 9: 22-40.

Wayne´s Word (by W.P. Armstrong):
! Bryophyte Photos.

Wikipedia, the free encyclopedia:
Also in German:

Wikipedia, the free encyclopedia:

! N. Wikström et al. (2009): Liverworts (Marchantiophyta). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life.
A version archived by Internet Archive Wayback Machine.
See also here.

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This index is compiled and maintained by Klaus-Peter Kelber, Würzburg,
Last updated November 11, 2020

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