Links for Palaeobotanists

Home / Introductions to both Fossil and Recent Plant Taxa / Bryophyta

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

N. Anantaprayoon et al. (2024): Evolution of the most species-rich family of simple thalloid liverworts (Aneuraceae): a time-calibrated perspective into its evolutionary history and diversification. In PDF,

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

V.A. Bakalin and S.S. Choi (2023): The Type Specimens of Plectocolea and Solenostoma (Marchantiophyta) in Some Japanese and European Herbaria. Open access, Plants, 12, 3935.

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).

! J. Bechteler et al. (2023): Comprehensive phylogenomic time tree of bryophytes reveals deep relationships and uncovers gene incongruences in the last 500 million years of diversification. Free access, American Journal of Botany,
"... The phylogeny resolves many relationships across the bryophytes, enabling us to resurrect five liverwort orders and recognize three more, and propose ten new orders of mosses. Most orders originated in the Jurassic and diversified in the Cretaceous or later ..."

Michael Becker: Moose (in German).
Still available via Internet Archive Wayback Machine.

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. 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.

! A.C. Bippus et al. (2022): The Role of Paleontological Data in Bryophyte Systematics. Abstract, Journal of Experimental Botany.
"... Paucity of the bryophyte fossil record, driven primarily by phenotypic (small plant size) and ecological constraints (patchy substrate-hugging populations), and incomplete exploration, results in many morphologically isolated, taxonomically ambiguous fossil taxa. Nevertheless, instances of exquisite preservation and pioneering studies demonstrate the feasibility of including bryophyte fossils in evolutionary inference. ..."

! A.C. Bippus (2022): Permineralized Mesozoic Moss Gametophytes and their Implications for Bryophyte Evolution. In PDF, Dissertation (abstract), Oregon State University.
See also here and there.

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.

! B. Bomfleur et al. (2023): Fossil mosses from the Early Cretaceous Catefica mesofossil flora, Portugal–a window into the Mesozoic history of Bryophytes. In PDF, Fossil Imprint, 79: 103–125.
See likewise here.
"... A diverse assemblage of mosses from the Early Cretaceous Catefica mesofossil flora, Portugal, is described based on fragments of charcoalified and lignitized gametophytes and a single spore capsule ..."

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

J.L. Bowman et al. (2022): The renaissance and enlightenment of Marchantia as a model system. Free access, The Plant Cell, 34: 3512–3542. Worth checking out:
Note fig. 1: Phylogenetic history of Marchantia plotted against the geologic timescale.
Figure 2. Life cycle of Marchantia polymorpha.

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:
What is Paleobotany?. Also worth checking out:
Plant Evolution & Paleobotany. An educational resource for students and teachers studying Earth's history, fossils, and evolution.
! Go to: Paleobotany Short-Course. Lecture notes.
Paleobotany Overview; Life moves to land.
Plant classification.
Rise of Seed Plants.
Rise of flowering plants.

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.

D.A. Callaghan et al. (2022): Long-term survival of bryophytes underground: an investigation of the diaspore bank of Physcomitrium eurystomum Sendtn. In PDF, Journal of Bryology, DOI: 10.1080/03736687.2022.2151857.
See also here.
"... Undisturbed soil cores of 40 cm depth were collected from Langmere, Norfolk, UK, and were split into investigated sediment layers of 1 cm depth.
[...] Viable diaspores of Physcomitrium eurystomum frequently occurred in sediment layers that were at least 100 years old and continued to occur in much lower layers that were probably several hundred years old.

! Southern Illinois University, Carbondale: Bryophytes.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

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.

P.M. Delaux and S. Schornack (2021): Plant evolution driven by interactions with symbiotic and pathogenic microbes. In PDF, Science, American Association for the Advancement of Science (AAAS), 371 (6531), pp.eaba6605. ff10.1126/science.aba6605ff. ffhal-03327916ff.
See also here.
"... Delaux and Schornack review how insights from a range of plant and algal genomes reveal sustained use through evolution of ancient gene modules as well as emergence of lineage-specific specializations. Mosses, liverworts, and hornworts have layered innovation onto existing pathways to build new microbial interactions. ..."

! P.M. Delaux et al. (2019): Reconstructing trait evolution in plant evo–devo studies. Free access, Current Biology, 29: R1110-R1118.
"... we summarize a subset of the different aspects of plant evolutionary biology, provide a guide for structuring comparative biology approaches and discuss the pitfalls that (plant) researchers should avoid when embarking on such studies ..."

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. (2021): Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions. Open access, Bryophyte Diversity and Evolution, 43.

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. Free access, Plant Syst. Evol., 299: 585-601.

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

B.J. Fletcher et al. (2005): Fossil bryophytes as recorders of ancient CO2 levels: Experimental evidence and a Cretaceous case study. Free access, Global Biogeochemical Cycles.

J.R. Flores et al. (2023): Dating the evolution of the complex thalloid liverworts (Marchantiopsida): total-evidence dating analysis supports a Late Silurian-Early Devonian origin and post-Mesozoic morphological stasis. In PDF, New Phytologist, doi: 10.1111/nph.19254.
See also here.
"... Phylogenetic analyses were performed on a combined dataset of 130 discrete characters and 11 molecular markers
[...] Total-evidence dating analyses support the radiation of Marchantiopsida during Late Silurian-Early Devonian (or Middle Ordovician when the outgroup is excluded) and that of Ricciaceae in the Middle Jurassic ..."

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).

E. Frangedakis et al. (2023): What can hornworts teach us? Open access, Front. Plant Sci., 14:1108027. doi: 10.3389/fpls.2023.1108027

A.O. Frolov et al. (2022): The first discovery of mosses (Bryopsida) in the Lower Jurassic of Eastern Siberia. Open access, Journal of Palaeosciences, 71.

Anthony G. Futcher, Columbia Union College, Maryland: Plant Diversity. A lot of facts about plant groups, fungi, plant-like protists, and monerans, including taxonomy, life cycles, general structure, and representative genera. Go to:
Division Bryophyta - Liverworts, Hornworts and Mosses.
Still available through the Internet Archive´s Wayback Machine.

! P. Gerrienne et al. (2022): Earliest Evidence of Land Plants in Brazil. In PDF, In: Iannuzzi, R., Rößler, R., Kunzmann, L. (eds.): Brazilian Paleofloras. Springer.
See also here.
Note. fig. 3: Suggested life cycle of an early vascular plant from the early Devonian Rhynie Chert.
Fig. 4b: Suggested reconstruction of Cooksonia paranensis.
Fig. 5: Suggested life cycle of Cooksonia paranensis.

! 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.

L. Han et al. (2021): Liverwort fossils from the Late Triassic of Baiyin City, Gansu Province, and their geological significance. In PDF, Palaeoworld. See also here.

B.J. Harris et al. (2021): Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants. bioRxiv, see also here.
Note figure 3: The timescale of land plant evolution.
"... Here we investigate the evolution of the land plants (embryophytes) and their two main lineages, the tracheophytes (vascular plants) and bryophytes (non-vascular plants).
[...] extant tracheophytes and bryophytes are both highly derived; as a result, understanding the origin of land plants requires tracing character evolution across the diversity of modern lineages.

B.J. Harris et al. (2020): Phylogenomic evidence for the monophyly of bryophytes and the reductive evolution of stomata: Free access, Current Biology, 30: 2001-2012.
"... Our analyses recover bryophyte monophyly and demonstrate that the guard cell toolkit is more ancient than has been appreciated previously.
[...] the first embryophytes possessed stomata that were more sophisticated than previously envisioned and that the stomata of bryophytes have undergone reductive evolution, including their complete loss from liverworts ..."

! 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, 165: 235-240.
See also here.

! 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, 40: 755–758.
See also here.

! M.S. Ignatov and E.V. Maslova (2021): Fossil mosses: What do they tell us about moss evolution? Open access, Bry. Div. Evo., 43.

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.

D.S. Kopylov et al. (2020): The Khasurty Fossil Insect Lagerstätte. In PDF, Paleontological Journal, 54: 1221–1394. See also here.
Worth checking out:
Starting on page 1350 (PDF page 130): Bryophyta and Marchantiophyta. Mosses and Liverworts (by Y.S. Mamontov and M.S. Ignatov).
Starting on page 1364 (PDF page 144): Trachaeophyta. Vascular Plants (by N.V. Bazhenova).

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).
"... Our results challenge the long-held notion that spores, and not vegetative propagules, are involved in long-distance dispersal. Asexual reproduction seems to play a major role in shaping the global distribution patterns of liverworts ..."

! 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. (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. Free access, Annals of botany, 109: 851–871.

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.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

S. McLoughlin and C. Strullu-Derrien (2015): Biota and palaeoenvironment of a high middle-latitude Late Triassic peat-forming ecosystem from Hopen, Svalbard archipelago. PDF file, in: Kear B.P. et al. (eds): Mesozoic Biotas of Scandinavia and its Arctic Territories. Geol. Soc. London Spec. Pub., 434: 87–112.
See also here.
Permineralized bryophyte sporangium!

S.A.M. McAdam et al. (2021): Stomata: the holey grail of plant evolution. In PDF, Am. J. Bot., 108: 366–371. See also here.

! 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!

R.L. Mitchell et al. (2023): Terrestrial surface stabilisation by modern analogues of the earliest land plants: A multi-dimensional imaging study. Open access, Geobiology.
Note figure 1: Summary chart highlighting the evolution of different CGC elements [cryptogamic ground covers] from contrasting molecular, phylogenetic and fossil dating methods, and schematic land plant phylogeny of modern terrestrial organisms, focussing on the bryophytes and specific liverwort genera.

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

J.V. Mundim et al. (2022): Small areas and small plants: Updates on Antarctic bryophytes. Free access, Acta Bot. Bras., 35.

! 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.

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.

Northern Ontario Plant Database (NOPD).
NOPD is a website that provides free public access to records of herbarium specimens housed in northern Ontario educational and government institutions. Go to:
Bryophyte Links.

! Oxford Bibliographies.
Oxford Bibliographies offers exclusive, authoritative research guides. Combining the best features of an annotated bibliography and a high-level encyclopedia, this cutting-edge resource directs researchers to the best available scholarship across a wide variety of subjects. Go to:
Fossils (by Kevin Boyce).
Evolution of Land Plants (by Charles C. Davis and Sarah Mathews).
Evolution of Fungi (by David Hibbett).
Bryophyte Ecology (by Heinjo During).

J. Patiño et al. (2022): Unveiling the nature of a miniature world: a horizon scan of fundamental questions in bryology. In PDF, Journal of Bryology, 44.
See also here.
"... comparative studies ofbryophytes across both taxonomic groups (including fossils) and spatial scales are necessary
[...] Past levels of bryophyte diversity [...] are probably underestimated, because the bryophyte fossil record, despite ongoing discoveries [...], remains scarce due to either the limited resistance of the plant body to decay or taphonomic biases and related issues ..."

! J. Patiño and A. Vanderpoorten (2018): Bryophyte Biogeography. In PDF, Critical Reviews in Plant Sciences, DOI: 10.1080/07352689.2018.1482444. See also here.
Note figure 2: Worldwide patterns of bryophyte hot spots of endemism.

! Phyto Images
Database hosted by the Cornell University Vascular Plant Herbarium. Software deveopment by Kevin C. Nixon and Jan De Laet).
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. Superbly done!

(Stefan A. Rensing University of Freiburg, Faculty of Chemistry and Pharmacy).
Plantcode scientists are are interested in the evolution of plants sensu lato - i.e., the photosynthetic eukaryotes. Go to:
List of publications.

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. Free access, Plant Physiology, 177: 788–797.

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. Free access, Am. J. Bot., 103: 652-662.

A.A. Santos et al. (2022): First evidence of Ricciaceae in the Jurassic of the Iberian Peninsula (Asturias, NW Spain): Ricciopsis asturicus sp. nov. Abstract, Botany Letters, 169: 557-567.

! 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. Free access, Am. J. Bot., 98: 352-369.

! 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).

M. Stech et al. (2021): Advances and challenges in bryophyte biology after 50 years of International Association of Bryologists. Free access, Bryophyte Diversity and Evolution, 43: 006–009.
See also here.

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.

D. Su et al. (2021): Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants. Free access, Molecular Biology and Evolution, 38: 3332–3344.
"... we estimate that land plants originated in the Precambrian (980–682 Ma), much older than widely recognized. Our study highlights the important contribution of molecular data when faced with contentious fossil evidence, and that fossil calibrations used in estimating the timescale of plant evolution require critical scrutiny. ..."

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.

Tarleton State University, (by A. Nelson (?)), Stephenville, Texas A&M University System:
Plant Evolution and Diversity. Powerpoint presentation.

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

W.A. Taylor and P.K. Strother (2024): Plant evolution: A tapetum is now effectively present in all land plant lineages. In PDF, Current Biology, 34: R146-R148. DOI: 10.1016/j.cub.2023.12.061.
See likewise here.
"... Three independent lines of evidence lead to the observation that the evolution of the plant spore occurred before the evolution of axial, upright plants ..."

! 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.

! L. VanAller 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.

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

K. van Zuijlen et al. (2023): Bryophytes of Europe Traits (BET) data set: A fundamental tool for ecological studies. Open access, Journal of Vegetation Science, 34. 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.
This expired link is now available through the Internet Archive´s Wayback Machine. Also worth checking out:
Northern Ontario Plant Database (NOPD).
NOPD is a website that provides free public access to records of herbarium specimens housed in northern Ontario educational and government institutions. Go to:
Bryophyte Links.

! 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.

! S. Woudenberg et al. (2022): Deep origin and gradual evolution of transporting tissues: Perspectives from across the land plants. In PDF, Plant Physiology.
See also here. Note figure 4: Summary of the early fossil record of transporting tissues.

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Last updated May 14, 2024

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