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! 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.

Michael Becker: Moose (in German).

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

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

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.

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.

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

! 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.

dmoz, the Open Directory Project:
Science: Biology: Flora and Fauna: Plantae:
Bryophyte. See also:
Earth Sciences: Paleontology: Paleobotany: Taxa.

dmoz: Science: Biology: Flora and Fauna: Plantae: Bryophyta.

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. (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.

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

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

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).

! 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.

Joannes A. Janssens, Minnesota Ground Water Association: Introduction to Bryology. Powerpoint presentation. See also here.

J.A. Janssens (2007): Description of Bryophyte Assemblages of Mesohabitats. In PDF. See also here.

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.

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: Sporophytes and gametophytes of Dicranaceae from the Santonian (Late Cretaceous) of Georgia, USA.

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

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?

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. (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. Snapshot taken by the Internet Archive´s Wayback Machine. Go to:
Plant anatomical characteristics.

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!

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!

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.

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.

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. In PDF, Amer. J. Botany, 91: 1557-1581.

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. 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. In PDF, 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.

! 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 (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:

! 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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Last updated March 03, 2017

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