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Permian Palaeobotany
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! Focussed on the Fossil Record@
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Progress in Palaeobotany and Palynology@
! Fungal Wood Decay: Evidence from the Fossil Record@
Abscission and Tissue Separation in Fossil and Extant Plants@
Stomatal Density@
Permineralized Plants and Petrified Forests@
! Chemotaxonomy and Chemometric Palaeobotany@

Carboniferous Palaeobotany

C. Álvarez-Vázquez and R.H. Wagner (2017): A revision of Annularia and Asterophyllites species from the lower Westphalian (Middle Pennsylvanian) of the Maritime Provinces of Canada. Abstract, Atlantic Geology, 53: 17-62. See also here (in PDF).

! J.M. Anderson et al. (1999): Patterns of Gondwana plant colonisation and diversification. In PDF, Journal of African Earth Sciences, 28: 145-167.
See also here.

P. Appleton et al. (2015; article starts on PDF page 21): Making the most of Brymbo’s plant fossils . Earth Heritage, 43.
Still available via Internet Archive Wayback Machine.
Note photograph on PDF page 22: sediment-filled casts of Calamites stems in growth position.

A.R. Bashforth et al. (2022): Taphonomic megabiases and the apparent rise of the dryland biome during the Pennsylvanian to Permian transition. Powerpoint presentation (pptx-extension), 11th European Palaeobotany and Palynology Conference (Stockholm, Sweden).

A. Bashforth et al. (2016): A Middle Pennsylvanian macrofloral assemblage from wetland deposits in Indiana (Illinois Basin): a taxonomic contribution with biostratigraphic, paleobiogeographic, and paleoecologic implications. In PDF, Journal of Paleontology, 90: 589–631.

A.R. Bashforth et al. (2016): Dryland vegetation from the Middle Pennsylvanian of Indiana (Illinois Basin): The dryland biome in glacioeustatic, paleobiogeographic, and paleoecologic context. Journal of Paleontology, 40: 785–814.

A.R. Bashforth et al. (2014): Paleoecology of Early Pennsylvanian vegetation on a seasonally dry tropical landscape (Tynemouth Creek Formation, New Brunswick, Canada). In PDF, Review of Palaeobotany and Palynology, 200: 229–263. See also here.
Note fig. 6, 7: Upright cordaitalean trees.
Fig. 8C, 8D: Upright Calamites axes.

A.R. Bashforth and W.A. DiMichele (2012): Permian Coal Forest offers a glimpse of late Paleozoic ecology. In PDF, PNAS, 109: 4717-4718.

A.R. Bashforth et al. (2010): Vegetation heterogeneity on a Late Pennsylvanian braided-river plain draining the Variscan Mountains, La Magdalena Coalfield, northwestern Spain. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology.

A.R. Bashforth (1999): Descriptive taxonomy, biostratigraphic correlation and paleoenvironmental reconstruction of an Upper Carboniferous macrofloral assemblage, Bay St. George Basin, Southwestern Newfoundland. Thesis, Memorial University of Newfoundland. See also here (in PDF).

R.M. Bateman et al. (2016): Stratigraphy, palaeoenvironments and palaeoecology of the Loch Humphrey Burn lagerstätte and other Mississippian palaeobotanical localities of the Kilpatrick Hills, southwest Scotland PeerJ, 4.

R.W. Baxendale (1979): Plant-bearing coprolites from North-American Pennsylvanian coal balls. PDF file.

D. J. Beerling et al.(1998): The influence of Carboniferous palaeoatmospheres on plant function: an experimental and modelling assessment. PDF file, Philosophical Transactions of the Royal Society B, 353, 131-140.

! J.H. Calder et al. (2006): A fossil lycopsid forest succession in the classic Joggins section of Nova Scotia: Paleoecology of a disturbance-prone Pennsylvanian wetland. Abstract, in: S.F. Greb and W.A. DiMichele (eds.): GSA Special Papers, Wetlands through Time, 399: 169-194. See also here (in PDF), and there (Google books).

C.J. Cleal et al. (2016): Spondylodendron pranabii — the dominant lycopsid of the late Mississippian vegetation of the Kashmir Himalaya. Abstract, Alcheringa: An Australasian Journal of Palaeontology, 40. See also here (in PDF).

! C.J. Cleal et al. (2015): Pennsylvanian fossil flora from the Velebit Mountains and Lika region (SW Croatia). In PDF, Bulletin of Geosciences, 90: 721-742.

! C.J. Cleal et al. (2012): Plant biodiversity changes in Carboniferous tropical wetlands. In PDF, Earth-Science Reviews, 114: 124-155.
See also here.
"... Using a combination of species richness, polycohort and constrained cluster analyses, the plant biodiversity of Pennsylvanian (late Carboniferous) tropical wetlands (“coalswamps”) has been investigated in five areas in Western Europe and eastern North America ..."

! C.J. Cleal et al. (2011): Pennsylvanian vegetation and climate in tropical Variscan Euramerica. In PDF, Episodes, 34.

C.J. Cleal & B. A. Thomas: A Provisional World List of Geosites for Palaeozoic Palaeobotany. Initiated by the IUGS to develop an inventory of globally important geological sites. GEOSITES provide a provisional list of candidate Palaeozoic palaeobotany sites. The results are summarized in 40 sites, which are intended to show the broad pattern of evolution in land floras from the middle Silurian to the end of the Permian.
Still available via Internet Archive Wayback Machine.
See also here.

V. Dernov (2019): Taphonomy and paleoecology of fauna and flora from deltaic sandstones of Mospinka Formation (Middle Carboniferous) of Donets Basin. In PDF, Geo & Bio, 18: 37–63.

W.A. DiMichele et al. (2017): Vegetational zonation in a swamp forest, Middle Pennsylvanian, Illinois Basin, U.S.A., indicates niche differentiation in a wetland plant community. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 487: 71–92. See also here.

W.A. DiMichele et al. (2017): Plant Fossils from the Pennsylvanian–Permian Transition in Western Pangea, Abo Pass, New Mexico. In PDF, Smithsonian Contributions to Paleobiology, 99.

! W.A. DiMichele (2014): Wetland-Dryland Vegetational Dynamics in the Pennsylvanian Ice Age Tropics. Int. J. Plant Sci., 175: 123-164. See also here (in PDF).
Large Sigillaria stump cast on PDF page 12 (fig. 8).
! Reconstructions of coal swamps and some dryland plant reconstructions with Cordaitalean trees Walchian conifers.

! W.A. DiMichele and H.J. Falcon-Lang (2011): Pennsylvanian "fossil forests" in growth position (T0 assemblages): origin, taphonomic bias and palaeoecological insights. PDF file, Journal of the Geological Society, London, 168: 585-605. See also here.
Note fig. 14 (PDF page 17), Animals using hollow Sigillarian stumps as refuges from fire.

William A. DiMichele et al. (2010): Cyclic changes in Pennsylvanian paleoclimate and effects on floristic dynamics in tropical Pangaea. PDF file, International Journal of Coal Geology, 83: 329-344. See also here.

! W.A. DiMichele et al. (2007): Ecological gradients within a Pennsylvanian mire forest. In PDF Geology, 35: 415–418.
See also here.
"... we report the discovery of a spectacular fossil forest preserved over -1000 ha
[...] The forest was abruptly drowned when fault movement dropped a segment of coastal mire below sea level. ..."

! W.A. DiMichele et al. (2006): From wetlands to wet spots: Environmental tracking and the fate of Carboniferous elements in Early Permian tropical floras. PDF file. In Greb, S.F., and DiMichele, W.A., Wetlands through time: Geological Society of America Special Paper 399, p. 223–248. See also here and there (Google books).

! W.A. DiMichele et al. (2004): An unusual Middle Pennsylvanian flora from the Blaine Formation (Pease River Group: Leonardian-Guadalupian Series) of King County, West Texas. Abstract, Journal of Paleontology, 78: 765-782.
See also here (in PDF).
Paper awarded with the "Winfried and Renate Remy Award 2005", The Botanical Society of America.

! W.A. DiMichele and T.L. Phillips (2002): The ecology of Paleozoic ferns. In PDF, Review of Palaeobotany and Palynology, 119: 143-159.
See also here.

William A. DiMichele et al. (2001): Response of Late Carboniferous and Early Permian plant communities to climate change. PDF file, Annual Review of Earth and Planetary Sciences, 29: 461-4871.

! W.A. DiMichele and T.L. Phillips (1996): Clades, ecological amplitudes, and ecomorphs: phylogenetic effects and persistence of primitive plant communities in the Pennsylvanian-age tropical wetlands. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 127: 83-105.
See also here.

M.P. Donovan et al. (2021): Atlas of Selected Kinney Quarry Plant Fossils, Late Pennsylvanian, Central New Mexico. Google books. PDF download available.
In: Lucas, S.G., DiMichele, W.A. and Allen, B.D., (eds.): Kinney Brick Quarry Lagerstätte. New Mexico Museum of Natural History and Science Bulletin, 84.

M.T. Dunn et al. (2012): Winslowia tuscumbiana gen. et sp. nov. (Chaloneriaceae): A Cormose, Heterosporous, Ligulate Lycopsid Reconstructed from the Inside Out from the Pride Mountain Formation (Late Mississippian/Serpukhovian) of Northern Alabama. Abstract, International Journal of Plant Sciences, 173: 96-111.

H.J. Falcon-Lang (2021): Climate–vegetation models bring fossil forests back to life. Free access, PNAS, 118.

H.J. Falcon-Lang et al. (2018): New insights on the stepwise collapse of the Carboniferous Coal Forests: Evidence from cyclothems and coniferopsid tree-stumps near the Desmoinesian–Missourian boundary in Peoria County, Illinois, USA. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 490: 375–392. See also here and there.

H.J. Falcon-Lang and W.A. DiMichele (2010): What happened to the coal forests during Pennsylvanian glacial phases? PDF file, Palaios, 25: 611-617. See also here.
Including a reconstruction of the Late Pennsylvanian ecosystem (fig 4).
"... plant assemblages in this paleoclimatic context suggests that coal forests dominated during humid interglacial phases, but were replaced by seasonally dry vegetation during glacial phases. After each glacial event, coal forests reassembled with largely the same species composition. ..."

! H.J. Falcon-Lang et al. (2006): The Pennsylvanian tropical biome reconstructed from the Joggins Formation of nova Scotia, Canada. In PDF, Journal of the Geological Society, London, 163: 561–576. See also here.

H.J. Falcon-Lang (2005): Adpressed tree-fern trunks from the Early Pennsylvanian Joggins Formation of Nova Scotia. In PDF, Atlantic Geology, 41: 169–172.

J. Galtier et al. (1992): Anatomically preserved conifer-like stems from the upper Carboniferous of England. In PDF, Proceedings of the Royal Society B: Biological sciences, 247. See also here.

! R.A. Gastaldo et al. (2020): The Coal Farms of the Late Paleozoic. In PDF. See also here.

R.A. Gastaldo et al. (2004): Erect forests are evidence for coseismic base-level changes in Pennsylvanian cyclothems of the Black Warrior Basin, USA. PDF file, in: J.C. Pashin and R.A. Gastaldo (eds): Sequence stratigraphy, paleoclimate, and tectonics of coal-bearing strata. AAPG Studies in Geology 51: 219-238.

! R.A. Gastaldo et al. (2004): Community heterogeneity of Early Pennsylvanian peat mires. Abstract.

S.F. Greb et al. (2006): Evolution and Importance of Wetlands in Earth History. PDF file, In: DiMichele, W.A., and Greb, S., eds., Wetlands Through Time: Geological Society of America, Special Publication, 399: 1-40.
Rhacophyton and Archaeopteris in a Devonian wetland as well as Pennsylvanian, Permian, Triassic and Cretaceous wetland plant reconstructions.
Note figure 1: Evolution of wetland types in the Silurian and Devonian.
See also here.
Still available through the Internet Archive´s Wayback Machine.

M. Grey and Z.V. Finkel (2011): The Joggins Fossil Cliffs UNESCO World Heritage site: a review of recent research. In PDF. Carboniferous forest reconstruction on page 192.

! A.J. J. Hetherington et al. (2016): Networks of highly branched stigmarian rootlets developed on the first giant trees. In PDF, PNAS, 113.

D. Hibbett et al. (2016): Climate, decay, and the death of the coal forests. In PDF, Current Biology, 26. See also here.

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.

K.R. Johnson (2007): Forests frozen in time. In PDF. Fig. 1 shows the reconstruction of a lycopsid forest.

! H. Kerp et al. (2007): Vegetationsbilder aus dem saarpfälzischen Permokarbon. PDF file, in German. In: Schindler, T, Heidtke, U.H.J. (eds.): Kohlesümpfe, Seen und Halbwüsten. Pollichia, Sonderveröffentlichung. See also here, and there (table of contents).

Hans Kerp, Palaeobotanical Research Group, Westfälische Wilhelms University, Münster: A History of Palaeozoic Forests. An introductory text with many helpful links directly related to the history of Palaeozoic forests. 7 chapters provide information about: The earliest land plants; Towards a tree-like growth habit; The earliest forests; The Carboniferous coal swamp forests; The floral change at the end of the Westphalian; Stefanian and Rotliegend floras; Is there a floral break in the Permian?
Now provided by the Internet Archive´s Wayback Machine.

! M. Krings et al. (2003): How Paleozoic vines and lianas got off the ground: on scrambling and climbing Carboniferous-early Permian pteridosperms. In PDF, The Botanical Review, 69: 204–224.
See also here.

! George Langford, "georgesbasement": Fossil Flora and Fauna of the Pennsylvanian Period, Will County, Illinois. Many fossil plant photographs, line drawings and reconstructions.
Links in the scientific names point to plates in Leo Lesquereux´s classic 1879 work, Atlas to the Coal Flora of Pennsylvania and of the Carboniferous Formation throughout the United States. See the Index to Fossil Flora, pp 1-85..
Collecting Fossil Plants and Animals in the Pennsylvanian Deposits of the Will County, Illinois Coal Measures The Field Notes of George Langford, Sr. in the Years 1937-1960. Prepared and organized by George Langford, Jr., 1973.
See also here.
These expired links is still available through the Internet Archive´s Wayback Machine.

M. Libertín et al. (2014): New sphenophyllaleans from the Pennsylvanian of the Czech Republic. In PDF, Review of Palaeobotany and Palynology, 200: 196-210.
See also here.

C.V. Looy et al. (2014): Evidence for coal forest refugia in the seasonally dry Pennsylvanian tropical lowlands of the Illinois Basin, USA. PeerJ., 2.

M. Lubienski (2013): Fossile Pflanzen aus dem Oberkarbon bei Albringhausen (Wetter, Ennepe-Ruhr-Kreis, Nordrhein-Westfalen). PDF file, in German. Jahrb. Bochumer Bot. Ver., 4: 9-30.

! L. Luthardt et al. (2021): Medullosan seed ferns of seasonally-dry habitats: old and new perspectives on enigmatic elements of Late Pennsylvanian–early Permian intramontane basinal vegetation. In PDF, Review of Palaeobotany and Palynology, 288.
See also here.
Note figure 1: Stratigraphy and fossil record of the Medullosales in the context of palaeogeographic and palaeoclimatic developments in the late Paleozoic.
Figure 2: Transverse sections of stem taxa of medullosans with information on their stratigraphy, (palaeo-) geographic origin, taphonomy and palaeo-environment.
Also of interest in this context:
Pflanzliche Botschaften aus der Urzeit (by Tamara Worzewski, November 08, 2022,, in German).

Eugene Marinus, Department of Biodiversity and Conservation Biology, University of the Western Cape: Ferns in the Carboniferous Period (Powerpoint presentatation).
Now recovered from the Internet Archive´s Wayback Machine.

C. Martín-Closas et al. (2018): New palaeobotanical data from Carboniferous Culm deposits constrain the age of the Variscan deformation in the eastern Pyrenees. Abstract, Geologica acta, 16: 107-123. See also here and there (in PDF).

S.V. Naugolnykh (2012): A new Carboniferous pteridosperm of Angaraland: Angaranthus victorii Naugolnykh, gen. et spec. nov.(Angaranthaceae, fam. nov., Callistophytales). In PDF, Wulfenia. See also here.
Note figure 9: Reconstructions of Gondwanotheca sibirica reproductive organs.

! M.P. Nelsen et al. (2016): Delayed fungal evolution did not cause the Paleozoic peak in coal production. Proceedings of the National Academy of Sciences, 113: 2442-2447. See also here.

R. Neregato and J. Hilton (2019): Reinvestigation of the Enigmatic Carboniferous Sphenophyte Strobilus Cheirostrobus Scott and Implications of In Situ Retusotriletes Spores. In PDF, Int. J. Plant Sci., 180: 811–833. See also here.

! S. Opluštil et al. (2022): Carboniferous macrofloral biostratigraphy: an overview. Abstract, Geological Society, London, Special Publications, 512: 813-863.

C.P. Osborne et al.(2004): Biophysical constraints on the origin of leaves inferred from the fossil record. PDF file, PNAS, 101: 10360-10362.
This expired link is available through the Internet Archive´s Wayback Machine.

G. Pacyna and D. Zdebska (2012): Carboniferous plants preserved within sideritic nodules - a remarkable state of preservation providing a wealth of information. In PDF, Acta Palaeobotanica, 52: 247-269.

Paläontologische Gesellschaft: Fossil des Jahres 2018. About Lepidodendron (in German).

Mary Parrish, Smithsonian National Museum of Natural History: Reconstructing a Carboniferous Peat Swamp.

H.W. Pfefferkorn et al. (2017): Impact of an icehouse climate interval on tropical vegetation and plant evolution. In PDF, Stratigraphy, 14: 365-376. See also here.

H.W. Pfefferkorn et al. (2001): Modern tropical analogs for Carboniferous standing forests: Comparison of extinct Mesocalamites with extant Montrichardia. Abstract, Historical Biology, 15.

! T.L. Phillips et al. (1985): Stratigraphic and interregional changes in Pennsylvanian coal-swamp vegetation: environmental inferences. In PDF, International Journal of Coal Geology, 5: 43-109.
See also here.

! T.L. Phillips et al. (1976): Fossil peat of the Illinois basin: a guide to the study of coal balls of Pennsylvanian age. In PDF, Geoscience education, 11.

E. Reeves et al. (2023): Historic palaeobotanical collection reveals in situ microspores and pollen from Early Carboniferous (Tournaisian) ovules from the Ballagan Formation of Scotland. In PDF, Review of Palaeobotany and Palynology, 308.
See also here.

! Sarda Sahney et al. (2010): Rainforest collapse triggered Carboniferous tetrapod diversification in Euramerica. PDF file, Geology, 38: 1079-1082. See also here, and there (abstract).

J.W. Schneider et al. (2010): Euramerican Late Pennsylvanian/Early Permian arthropleurid/tetrapod associations - implications for the habitat and paleobiology of the largest terrestrial arthropod. PDF file, in: Lucas, S.G., Schneider, J.W. and Spielmann, J.A., (eds.): Carboniferous-Permian transition in Canon del Cobre, northern New Mexico: New Mexico Museum of Natural History and Science, Bulletin 49: 49-70.

Andrew C. Scott et al. (2009): Scanning Electron Microscopy and Synchrotron Radiation X-Ray Tomographic Microscopy of 330 Million Year Old Charcoalified Seed Fern Fertile Organs. PDF file, Microsc. Microanal., 15: 166-173.
See figure 4, SEM of charcoalified pteridosperm ovule from the mid-Mississippian (Carboniferous). See also here.

B. Slater (2011): Fossil focus: Coal swamps. n PDF, Palaeontology Online. See also here.

! G.W. Stull et al. (2012): Palaeoecology of Macroneuropteris scheuchzeri, and its implications for resolving the paradox of "xeromorphic" plants in Pennsylvanian wetlands. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 331-332: 162-176.
See also here.

C. Strullu-Derrien et al. (2023): The Carboniferous (Serpukhovian) macroflora from the “Coteaux du Pont Barré”, Beaulieu-sur- Layon (Maine-et-Loire), South of the Armorican Massif, France. Open access, Botany Letters.
See also here (in PDF).

! B.A. Thomas and C.J. Cleal (2022): A reassessment of the leafy shoots of Pennsylvanian-age arborescent lycopods. Open acces, Botany Letters, DOI: 10.1080/23818107.2022.2101517.
See also here.
Note figure 1: Reconstruction of arborescent lycopsids of the Pennsylvanian-age palaeotropical coal swamps of Euramerica.

! B.A. Thomas et al. (2019): The distribution of plant fossils and their palaeoecology in Duckmantian (Bashkirian, Lower Pennsylvanian) strata at Brymbo, North Wales, UK. Open access, Geological Journal.
Note figure 3b: Stigmaria trunk in situ.
Note figure 17: Calamites stems and pith casts.

! B.A. Thomas and L.J. Seyfullah (2015): Stigmaria Brongniart: a new specimen from Duckmantian (Lower Pennsylvanian) Brymbo (Wrexham, North Wales) together with a review of known casts and how they were preserved. Abstract, Geological Magazine, 152: 858–870. See also here (in PDF).

A. Tosal et al. (2022): Plant taphonomy and palaeoecology of Pennsylvanian wetlands from the Erillcastell Basin of the eastern Pyrenees, Catalonia, Spain. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 605.
See also here.
"... A specimen of C. undulatus (50 cm long and 5 cm wide) was found charred and in an upright position within a pyroclastic bed intercalated in these shales ..."
Note figure 6; Plant taphonomic features. See especially:
Figure 6C: Charred Calamites undulatus stem crossing an ignimbrite deposit.

Susan Trulove, Virginia Tech: Ancient climate record preserved in prehistoric plants. Ancestor of modern trees preserves record of ancient climate change. About Devonian/Carboniferous growth rings.

! J.P. Wilson et al. (2017): Dynamic Carboniferous tropical forests: new views of plant function and potential for physiological forcing of climate. In PDF, New Phytologist, 215: 1333–1353. See also here.
! Figure 2 shows the fungal evolution and abundance of coal basin sediments over the Phanerozoic.

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Last updated March 17, 2023