Links for Palaeobotanists

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

Cyanobacteria and Stromatolites
Seed Plants in General
! Taxonomy and Plant Classification Databases@
! Living Fossils@
Plant Photographs@
Image Collections@
Picture Search@

Algae (previously GuruNet), New York City & Jerusalem: Coccolithophorida.
Provided by the Internet Archive´s Wayback Machine.

! Stanley M. Awramik, Department of Earth Science, University of California Santa Barbara:
The Record of Life on the Early Earth. Lecture notes, Powerpoint presentation.

AYMA Agua y Medio Ambiente, Sevilla: Atlas of Microorganisms. This expired link is available through the Internet Archive´s Wayback Machine.
Go to: Algae.

M.E. Barkworth et al. (2016): Report of the Special Committee on Registration of Algal and Plant Names (including fossils). In PDF, Taxon, 65: 670-672.

! B. Becker and B. Marin (2009): Streptophyte algae and the origin of embryophytes. In PDF, Annals of Botany, 103: 999–1004. See also here.
Note fig. 2: Diversification of green plants (Viridiplantae) and colonization of terrestrial habitats by streptophyte algae.

S. Bengtson et al. (2017): Three-dimensional preservation of cellular and subcellular structures suggests 1.6 billion-year-old crown-group red algae. Open Access, PLoS Biol., 15: e2000735.

! M. Berbee et al. (2020). Genomic and fossil windows into the secret lives of the most ancient fungi. In PDF, Nature Reviews Microbiology, 18: 717-730. 10.1038/s41579-020-0426-8.
See also here.
"... Inferences can be drawn from evolutionary analysis by comparing the genes and genomes of fungi with the biochemistry and development of their plant and algal hosts. We then contrast this emerging picture against evidence from the fossil record to develop a new, integrated perspective on the origin and early evolution of fungi ..."

The University of California Museum of Paleontology, Berkeley:
Introduction to the "Green Algae".

The University of California Museum of Paleontology, Berkeley: The Lake Winnipeg Algal Flora. Late Ordovician algae. See also:
Fossil Plants. Images from the UCMP collections.

Museum of Paleontology, University of California, Berkeley: "Green Algae": Systematics, Part 2, The Charophytes.

B. Bomfleur et al. (2010): Thalloid organisms and the fossil record - New perspectives from the Transantarctic Mountains. PDF file, Plant Signal Behav., 5: 293-295.

George Booth, The Fish Information Service: Algae. Easy to understand information.

! A.M.C. Bowles et al. (2023): The origin and early evolution of plants. Open access, Trends in Plant Science, 28.
Note figure 2: Phylogeny of early plant evolution with a selection of available genomic resources.
Figure 3: Fossils of possible and probable early archaeplastids.
! Figure 4: Summary of molecular estimates for the timescale of archaeplastid evolution.
"... Molecular clock analyses estimate that Streptophyta and Viridiplantae emerged in the late Mesoproterozoic to late Neoproterozoic, whereas Archaeplastida emerged in the late-mid Palaeoproterozoic ..."

Department of Biological Sciences, Bowling Green State University, Bowling Green, OH:
! Image Archive. Digitized diatom images. Excellent!
Freshwater Algae Links.
Marine Algae Links.

! J.J. Brocks et al. (2023): Lost world of complex life and the late rise of the eukaryotic crown. In PDF, Nature, See also here.
Note figure 1: Geological time chart comparing the molecular fossil, microfossil and phylogenetic records of early eukaryote evolution.

J.J. Brocks et al. (2017): The rise of algae in Cryogenian oceans and the emergence of animals. Abstract, Nature, 548: 578–581. See also here.

R.C. Brown and B.E. Lemmon (2011): Spores before sporophytes: hypothesizing the origin of sporogenesis at the algal-plant transition. In PDF, New Phytologist, 190: 875-881.

Mark Buchheim, Department of Biological Science, University of Tulsa, OK: deepestgreen. A website for the study of green algal diversity - a coordinated research effort for the phylogenetic investigation of the Chlorophyta. The Chlorophyta is one of two branches of the Viridiplantae (Green Plant) lineage. The Chlorophyta includes extant members that possess a fossil record that extends back into the Precambrian.

I.I. Bucur et al. (2020): Upper Triassic calcareous algae from the Panthalassa Ocean. Free access, Rivista Italiana di Paleontologia e stratigrafia, 126: 499-540.

Benjamin Burger, Utah State University, Vernal, Utah:
Why study fossil plants?
Invertebrate Paleontology and Paleobotany.
How did plants colonize the land, based on the fossil record?
How did the first seed plants (the Gymnosperms) evolve?
How did gymnosperms diversify during the early Mesozoic to become a modern dominate plant group?
How good is the fossil record of Cycads?
What is the significance of the fossil record of Ginkgo?
What is the fossil record of Horsetails?
! Fossil Algae.
What is an Angiosperm?
Video lectures.

! E.M. Carlisle et al. (2021): Experimental taphonomy of organelles and the fossil record of early eukaryote evolution. Open access, Science Advances, 7. DOI: 10.1126/sciadv.abe9487 See also here (in PDF).

Charophytes. A journal dedicated to the promotion of research and communication about charophytes.

! Michael Clayton, Department of Botany, University of Wisconsin, Madison: Instructional Technology (BotIT). Some image collections. Excellent! Go to:

N.R. Cúneo et al. (2014): Late Cretaceous Aquatic Plant World in Patagonia, Argentina. Open access, PLoS ONE, 9: e104749.

O. De Clerck et al. (2012): Diversity and Evolution of Algae: Primary Endosymbiosis. In PDF, Advances in Botanical Research, 64. See also here.
This expired link is available through the Internet Archive´s Wayback Machine.

! 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.F. Delwiche and E.D. Cooper (2015): The Evolutionary Origin of a Terrestrial Flora. Abstract, Current Biology. Please take notice:
! From algae to land plants (and vice versa). Did some freshwater algae descend from a terrestrial ancestor? In PDF.

The Delwiche Lab, Molecular Systematics: Charophycean Green Algae.

J. de Vries et al. (2018): Embryophyte stress signaling evolved in the algal progenitors of land plants. In PDF, PNAS, 115. See also here (abstract), and there (in German).

Emanuele Di Lorenzo, Georgia Institute of Technology, Atlanta, Georgia:
Early Earth and the Origins of Life.
Powerpoint presentation.

Earth Science Picture of the Day (EPOD). A service of USRA, sponsored by NASA Goddard. EPOD will collect and archive photos, imagery, graphics, and artwork with short explanatory captions and links exemplifying features within the Earth system. Browse EPODs by Related Fields, such as Coccolithophore bloom in the Celtic Sea.
These expired links are now available through the Internet Archive´s Wayback Machine.

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 Chlorophyta - Green Algae.
Still available through the Internet Archive´s Wayback Machine.

Janus Goulding (Derek Keats, Botany Department at the University of the Western Cape, Bellville (Cape Town) South Africa): Classification of Algae: where do they fit the broad scheme of things?
Snapshot taken by the Internet Archive´s Wayback Machine.

L.E. Graham (2019): Digging deeper: why we need more Proterozoic algal fossils and how to get them. Free access, Journal of phycology, 55: 1–6.

! L.E. Graham and L. W. Wilcox, Department of Botany, University of Wisconsin-Madison: Algae. A textbook (published by Prentice Hall), brief contents, including some pictures.
Still available via Internet Archive Wayback Machine.

Michael Guiry, Martin Ryan Institute, National University of Ireland, Galway, Ireland: What are Algae? A basic guide to algae, including freshwater, terrestrial and marine algae (seaweeds). See also:
The Seaweed Site. This site is a source of general information on all aspects of marine algae.

M.D. Guiry and E. Nic Dhonncha (project currently financed from the NUI, Galway, Environmental Change Institute funded by the Higher Education Authority (PRTLI Cycle II) in Ireland for 2000-2002): AlgaeBase. AlgaeBase is a dynamic, searchable database that stores information on the scientific names of seaweeds (including seagrasses), particularly taxonomic and distributional details. More than 20,000 seaweed and seagrass names are listed at present, but more are added each day. The data are being updated continually.

Till Hanebuth, Bastian Roters and Karl-Heinz Baumann, FB 05 Research group Sedimentology and Palaeoceanography, Bremen University, Germany: What are coccolithophores?

J. Hill and K. Davis, Geology Rocks: The Use Of Diatoms As Palaeoenvironmental Indicators.

Patrick Honecker, University of Cologne: Ancestors of land plants revealed.
Still available through the Internet Archive´s Wayback Machine.

! International "Fossil Algae" Association (IFAA). The IFAA is a non-profit organization interested in promoting the study of fossil algae, e.g. taxonomy, morphology, biology, biostratigraphy, palaeo-ecology and mineralization. Go to the archives. Reports, collections, reprints.

International Research Group on Charophytes (IRGC). The aims of IRGC are to promote and coordinate international cooperation in charophyte research, including living and fossil material of all geological periods, and to encourage the integration and synthesis of all aspects of both botanical and paleobotanical study of charophytes. Visit the IRGC photoalbum.

! Derek Keats, Botany Department at the University of the Western Cape, Bellville (Cape Town) South Africa: Introduction to algae (A virtual slide show), and The World of Algae.

! E. Kustatscher et al. (2022): A whole-plant specimen of the marine macroalga Pterigophycos from the Eocene of Bolca (Veneto, N-Italy). Open access, Fossil Imprint, 78: 145–156.
Note text-figure 5: Reconstruction drawing of Pterigophycos sp. thallus growing on a rock surface.

! M. Krings, LMU München: Unkalzifizierte fossile Makroalgen. Scientific project report (in German).

Landelijk Informatiecentrum voor Kranswieren (LIK), The Netherlands: Drawings and photos of recent charophyte species in the Netherlands (in Dutch).

! F. Leliaert et al. (2012): Phylogeny and Molecular Evolution of the Green Algae. PDF file, Critical Reviews in Plant Sciences, 31: 1-46.
See also here.

! F. Leliaert et al. (2011): Into the deep: new discoveries at the base of the green plant phylogeny. PDF file, BioEssays. 33: 683-692. See also here.
! Note figure 1: Phylogenetic relationships among the main lineages of green plants.
"... A schism early in their evolution gave rise to two major lineages, one of which diversified in the world’s oceans and gave rise to a large diversity of marine and freshwater green algae (Chlorophyta) while the other gave rise to a diverse array of freshwater green algae and the land plants (Streptophyta) ..."

! Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima. Go to:
Plant anatomical characteristics.

Ruta B. Limaye et al. (2007): Non-pollen palynomorphs as potential palaeoenvironmental indicators in the Late Quaternary sediments of the west coast of India. PDF file, CURRENT SCIENCE, VOL. 92, NO. 10.

! S.G. Lucas (2018): Permian-Triassic Charophytes: Distribution, Biostratigraphy and Biotic Events. Journal of Earth Science, 29: 778–793. See also here, and there (in PDF).

X. Ma et al. (2023): A reinvestigation of multiple independent evolution and Triassic-Jurassic origin of multicellular Volvocine algae. Open access, Genome Biology and Evolution, evad142,
"... The volvocine algae, a unique clade of chlorophytes with diverse cell morphology, provide an appealing model for investigating the evolution of multicellularity and development
[...] the dating analyses indicate that the volvocine algae occurred during the Cryogenian to Ediacaran (696.6–551.1 Ma), and multicellularity in the volvocine algae originated from the Triassic to Jurassic ..."

K.M. Maloney et al. (2022): Preservation of early Tonian macroalgal fossils from the Dolores Creek Formation, Yukon. Open access, Scientific Reports, 12.

! W.F. Martin and J.F. Allen (2018): An algal greening of land. Free access, Cell, 174: 256-258. See also here.
Note figure 1: Streptophyte Algae and the Rise of Atmospheric Oxygen.

C. Martín-Closas et al. (2009): Triassic charophytes from Slovenia: palaeogeographic implications. Abstract, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 252.

C. Martín-Closas (2003): The fossil record and evolution of freshwater plants: a review. PDF file, Geologica Acta, 1: 315-338.

Martin C. Mathes, College of William and Mary, Williamsburg, VA: General Botany. This course is designed to give the students a broad background in the traditional subject matter of botany. This includes topics on organisms in the plant kingdom as well as organisms not in the plant kingdom but which affect the growth ecology or evolution of plants (e.g., selected bacteria, fungi, and selected protists).

! C. Mays et al. (2021): Permian–Triassic non-marine algae of Gondwana—Distributions, natural affinities and ecological implications. Open access, Earth-Science Reviews, 212. See also here.

! R.M. McCourt et al. (2023): Green land: Multiple perspectives on green algal evolution and the earliest land plants. In PDF, American Journal of Botany 110. See also here (Free to read).
! Note figure 1: Green plant diversification in the context of the fossil record.
"... Green plants, broadly defined as green algae and the land plants (together, Viridiplantae), constitute the primary eukaryotic lineage that successfully colonized Earth's emergent landscape.
[...] We present the process not as a step-by-step advancement from primitive green cells to an inevitable success of embryophytes, but rather as a process of adaptations and exaptations that allowed multiple clades of green plants ..."

Richard M. McCourt et al. (2004): Charophyte algae and land plant origins. Abstract, Trends in Ecology and Evolution, 19.

! L.K. Medlin (2009): Haptophyte algae (Haptophyta), and Diatoms (Bacillariophyta). PDF files, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

L. Miao et al. (2024): 1.63-billion-year-old multicellular eukaryotes from the Chuanlinggou Formation in North China Science Advances, 10. DOI: 10.1126/sciadv.adk3208. See also here.
! Note figure 8: Overview of early evolution of the Eukarya along with fossil records.
"... we report cellularly preserved multicellular microfossils (Qingshania magnifica) from the ~1635-million-year-old Chuanlinggou Formation, North China. The fossils consist of large uniseriate, unbranched filaments with cell diameters up to 190 micrometers; spheroidal structures, possibly spores, occur within some cells ..."

M. Moczydlowska et al. (2011): Proterozoic phytoplankton and timing of chlorophyte algae origins. Open access, Palaeontology, 54: 721–733.

G.L. Mullins et al, Department of Geology, University of Leicester: The phytoPal project. About Proterozoic and Palaeozoic phytoplankton (fossil cysts of acritarchs, the phycomata of prasinophyte algae and very rare zygotes of zygnematalean algae).

! The Nannotax website (created by J.R. Young, P.R. Bown and J.A. Lees, International Nannoplankton Association). This website aims to provide an authoritative guide to the biodiversity and taxonomy of coccolithophores - a beautiful group of microscopic planktonic algae with an abundant fossil record. It is both a working tool for specialists and a reference source for anyone looking for information on coccolithophores.
Website awarded with the Golden Trilobite 2014.

! M.P. Nelsen et al. (2020): No support for the emergence of lichens prior to the evolution of vascular plants. In PDF, Gebiology, 18: 3-13. See also here.
! Note figure 2: Crown age estimates for LFF [lichenforming fungi] and putative origins of LFA [lichenforming algae].
"... As unambiguous fossil data are lacking to demonstrate the presence of lichens prior to vascular plants, we utilize an alternate approach to assess their historic presence in early terrestrial ecosystems. Here, we analyze new time-calibrated phylogenies of ascomycete fungi and chlorophytan algae
[...] Coupled with the absence of unambiguous fossil data, our work finds no support for lichens having mediated global change during the Neoproterozoic-early Paleozoic prior to vascular plants..."

K.J. Niklas and B.H. Tiffney (2022): Viridiplantae Body Plans Viewed Through the Lens of the Fossil Record and Molecular Biology. Open access, Integrative and Comparative Biology,
"... A review of the fossil record coupled with insights gained from molecular and developmental biology reveal a series of body plan transformations that gave rise to the first land plants. Across diverse algal clades, including the green algae and their descendants, the plant body plan underwent a unicellular -- colonial -- simple multicellular -- complex multicellular transformation series. ..."
Note figure 4: Scenarios for the evolution of the first land plant sporophyte resulting from delayed zygotic meiosis.

! K.J. Niklas and U. Kutschera (2010): The evolution of the land plant life cycle. Free access, New Phytologist, 185: 27-41.

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.

Charles J. O´Kelly and and Tim Littlejohn: Peridinium. The Protist Image Database.

Petralga. The PETRALGA (PErmian & TRiassic ALGAe) Project was initiated in order to provide useful palaeontological tools for both Scientific Institutions and Industry.

Z.A. Popper et al. (2011): Evolution and Diversity of Plant Cell Walls: From Algae to Flowering Plants. In PDF, Annu. Rev. Plant Biol., 62: 567-590.

Z.A. Popper and M.G. Tuohy (2010): Beyond the Green: Understanding the Evolutionary Puzzle of Plant and Algal Cell Walls. PDF file, Plant Physiology, 153: 373-383.

S.M. Porter (2004): The fossil record of early eukaryotic diversification. In PDF, Paleontological Society Papers, 10: 35-50.
Still available via Internet Archive Wayback Machine.
See also here.
Note figure 1: A current view of eukaryote phylogeny, based on a consensus of molecular and ultrastructural data.

Protist Information Server, Japan (supported by the "Soken-Taxa" project "Construction of Biological Image Databases" at The Graduate University for Advanced Studies, and by the "Bio-Resource" project "Fundamental research and development for databasing and networking culture collection information" at Japan Science and Technology Corporation). This server is providing 59932 images of protists (575+ genera, 2492+ species, 11645 samples) and 1294 movie clips as research and educational resources. See also: The Protist Movie Database.

C. Puginier et al. (2021): Plant–microbe interactions that have impacted plant terrestrializations. Free access, Plant Physiology.
Note figure 1: 1 Phylogenetic tree of the Viridiplantae. showing the evolution of the AMS [arbuscular mycorrhizal symbiosis], the putative evolutions of lichens and clades that contain LFA [lichen forming algae] and terrestrial species.
Figure 3: Lichens and their tolerance against terrestrial-related constraints.

Michael W. Rasser, Institute of Paleontology, Vienna: Fossil Coralline Algae.

S. Ratti et al. (2011): Did Sulfate Availability Facilitate the Evolutionary Expansion of Chlorophyll a+c Phytoplankton in the Oceans? In PDF, Geobiology 9, no. 4: 301–312. See also here (abstract).

P.K. Raval et al. (2023): A molecular atlas of plastid and mitochondrial adaptations across the evolution from chlorophyte algae to angiosperms. Free access, bioRxiv, doi:
"... Algae and plants carry two organelles of endosymbiotic origin that have been co-evolving in their host cells for more than a billion years. The biology of plastids and mitochondria can differ significantly across major lineages ..."

! J.A. Raven et al. (2012): Algal evolution in relation to atmospheric CO2: carboxylases, carbon-concentrating mechanisms and carbon oxidation cycles. Free access, Phil. Trans. R. Soc. B, 367: 493–507.

S.A. Rensing (2018): Great moments in evolution: the conquest of land by plants. Abstract, Current opinion in plant biology, 2018
"... Most probably, filamentous freshwater algae adapted to aerial conditions and eventually conquered land.
[...] In the past few years, the ever increasing availability of genomic and transcriptomic data of organisms representing the earliest common ancestors of the plant tree of life has much informed our understanding of the conquest of land by plants ..."

W.B. Sanders (2023): Is lichen symbiont mutualism a myth? Open access, BioScience, 73: 623–634.
See likewise here (in PDF).
Note figure 3: Two symbioses on intertidal rocks compared with respect to the lichen concept.
Figure 4: Two symbioses involving fungi of the Verrucariaceae (Ascomycota) and green algae of the Prasiolaceae compared with respect to the lichen concept.

P.A. Siver (2020): Remarkably preserved cysts of the extinct synurophyte, Mallomonas ampla, uncovered from a 48 Ma freshwater Eocene lake. In PDF, Scientific Reports, 10: 5204.

Department of Botany, Smithsonian Institution, Washington, DC: Algae Research. Go to: Classification, and Algae Links.

Geology Collection, Southampton Oceanography Centre (SOES), University of Southampton: SOES Geology Collection / Fossils / Algae and Stromatoporoids.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Fossil plant images from the oldest land plants. Go to: Prototaxites.

D. Su et al. (2022): Large-scale phylogenomic analyses reveal the monophyly of bryophytes and neoproterozoic origin of land plants Open access, Molecular Biology and Evolution, 38: 3332–3344.
! Note figure 1: The concatenation species tree of land plants and their algal relatives.
! Figure 2: The coalescent species tree of land plants and their algal relatives.
"... We found that studies favoring a Neoproterozoic origin of land plants (980–682 Ma) are informed more by molecular data whereas those favoring a Phanerozoic origin (518–500 Ma) are informed more by fossil constraints. Our divergence time analyses highlighted the important contribution of the molecular data (time-dependent molecular change) when faced with contentious fossil evidence.
[..] A careful integration of fossil and molecular evidence will revolutionize our understanding of how land plants evolved.

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
! BOT335 Lecture Schedule. Some interesting chapters in terms of palaeobotany, e.g.
The First Vascular Land Plants;
Carboniferous Forests;
Arborescent Lycopods;
Psaronius: a Carboniferous tree-fern;
Carboniferous Horsetails;
Carboniferous Seed Ferns;
The Evolution of Conifers;
Cycadophytes, the True Cycads;
Mesozoic Cycadeoids;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

! T.N. Taylor and M. Krings (2005): Fossil microorganisms and land plants: Associations and interactions. PDF file, Symbiosis, 40: 119-135.
This expired link is now available through the Internet Archive´s Wayback Machine.
See also here.

Teacher Certification:
The Algae Gallery. Including some links to other algae websites.
Still available through the Internet Archive´s

Bernard Teyssèdre, Paris: Are the green algae (phylum Viridiplantae) two billion years old? Carnets de Géologie / Notebooks on Geology: Article 2006/03.
See likewise here.

Sebastian Trapp, Bremen University: Charophytes in man-made lakes in Bremen, Germany.
This expired link is available through the Internet Archive´s Wayback Machine.

L.M. van Maldegem et al. (2019): Bisnorgammacerane traces predatory pressure and the persistent rise of algal ecosystems after Snowball Earth. Open access, nature communications.

A. Vicente et al. (2024): A bioprovince for the Barremian–Aptian charophytes of the Central Tethyan Archipelago. Free access, Cretaceous Research, 154.
See also here (in PDF):

! C.H. Wellmann et al. (2023). Terrestrialization in the Ordovician. Open access, Geological Society, London, Special Publications, 532.
"... This contribution reviews the evidence for terrestrial organisms during the Ordovician (microbial, land plant, fungal, animal)
[...] We conclude that the Ordovician was a critical period during the terrestrialization of planet Earth that witnessed the transition from a microbial terrestrial biota to one dominated by a vegetation of the most basal land plants. ..."

C.H. Wellman et al. (2019): Filamentous green algae from the Early Devonian Rhynie chert. Free access, PalZ.

Biology Department, Western Washington University, Bellingham, Washington:
! Protista - Algae. Powerpoint presentation. See also here, or there.

Wikipedia, the free encyclopedia.
Category:Algae classes.
Category:Green algae classes.
Category:Red algae classes.

Wikipedia, the free encyclopedia (in German).

Sabina Wodniok et al. (2011): Origin of land plants: Do conjugating green algae. PDF file, BMC Evolutionary Biology, 11: 104.

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