Home / Articles in 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), 11^th 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, Paleontology, 22: 537–548.
The link is to a version archived by the Internet Archive´s Wayback Machine.
See also here.

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.

N. Boyarina (2023): The Late Pennsylvanian vegetation of the Donets Basin, Ukraine: Syntaxonomy of plant communities. In PDF, GEO&BIO, 25: 64–98.

MSc Palaeobiology Students, Department of Earth Sciences, University of Bristol, (the author´s name appears on the title page for each section):
Fossil Lagerstätten. A catalogue of sites of exceptional fossil preservation. Go to:
Mazon Creek.
Websites still available via Internet Archive Wayback Machine.

! 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).

B. Cascales-Miñana et al. (2025): The taxonomy of fossil lyginopteridalean fronds Eusphenopteris neuropteroides and their importance for understanding late Carboniferous floristic changes. Open access, Taxon. https://doi.org/10.1002/tax.13317.

M.E. Chrpa et al. (2023): A marine origin of coal balls in the Midland and Illinois basins, USA. Open access, Communications Earth & Environment, 4.
"... Despite their importance to paleobotany, the salinity of coal-ball peat remains controversial. Pennsylvanian coal balls from the Midland and Illinois basins contain echinoderms and early high-magnesium calcite cement
[...] Coal balls likely formed in the marine-freshwater mixing zone ..."

! C.J. Cleal and B.A. Thomas (2023): Taxonomy and nomenclature of Sphenopteris and allied fossil-genera of Carboniferous seed-plant fronds. Free access, Taxon, 72: 862–879.
Note figure 10: Taxonomy and nomenclature of Sphenopteris and allied fossil-genera of Carboniferous seed-plant fronds.
"... Eight fossil-genera of lyginopteridalean fronds are now recognised (Sphenopteris, Calymmotheca, Eusphenopteris, Karinopteris, Mariopteris, Palmatopteris, Spathulopteris, Sphenopteridium) ..."

! C.J. Cleal and B.A. Thomas (2023): Taxonomy and nomenclature of Sphenopteris and allied fossil-genera of Carboniferous seed-plant fronds. Open access, Taxon, 72: 717-964.

! C.J. Cleal (2022): The Craigleith Tree (“Pitys withamii Tree”): Morphology, taxonomy, preservation and ecological context. In PDF.
Note figures on PDF page 6: Reconstruction of tree bearing the Pitys primaeva trunk (the Tweed Mill Tree) proposed by Retallack & Dilcher (1988).

C.J. Cleal (2022): The Craigleith Tree. In PDF.
"... The Craigleith Tree (Pitys withamii Tree) was a species of early seed plant, belonging to the general group known as the hydrasperman pteridosperms
[...] which indicate an early Asbian / late Visean age
[...] The trees were at least 20 m tall, with a trunk up to 1 m wide at the base, and were the tallest known woody trees growing anywhere in the world at this time.

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.

P.J. Coorough Burke et al. (2024): Mazon Creek Fossils Brought to You by Coal, Concretions, and Collectors. Abstract, Geological Society, London, Special Publications, 543.
"... The Mazon Creek biota includes over 465 animal and 350 plant species representing more than 100 orders, which is attributed to the preservation of organisms from multiple habitats and the large number of specimens collected. That phenomenon was made possible by coal extraction bringing concretions to the surface and highly motivated amateur collectors pursuing them ..."

M.P. D'Antonio and F. Herrera (2024): New Evidence of Unequal Branching in Stigmaria ficoides (Lycopsida). Open access, International Journal of Plant Sciences, 185.
"... Stigmaria ficoides is the main form species of rooting organ for late Paleozoic arborescent lycopsids in the families Diaphorodendraceae and Lepidodendraceae.
[...] we report two S. ficoides specimens based on the presence of the diagnostic rootlet scar pattern ..."

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.

D.L. Dilcher et al. (2005): Fossil Plants from the Union Chapel Mine, Alabama. PDF file, from: Buta, R.J., Rindsberg, A.K., and Kopaska-Merkel, D.C., eds., 2005, Pennsylvanian Footprints in the Black Warrior Basin of Alabama. Alabama Paleontological Society Monograph no. 1.
Images of Lepidophloios, Lepidodendron, Lepidostrobus, Lepidostrobophyllum, Lepidophylloides, Calamites, Calamostachys, Asterophyllites charaeformis, phenopteris, Neuralethopteris, Trigonocarpus ampulliforme, Whittleseya elegans.
Still available via Internet Archive Wayback Machine.

W.A. DiMichele et al. (2023): PALEOBOTANY OF THE CORINTH COAL BED, UPPER PENNSYLVANIAN, SOUTHERN ILLINOIS. In PDF, in: Lucas et al. (eds.), Fossil Record 9. New Mexico Museum of Natural History and Science Bulletin 94.

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. (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.

! W.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: 461-487.
See also here.

! 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. (2024): To rush into the secret house of death: The fate of a Tournaisian plant Geology, 20.
"... Tournaisian-age failure of marginal lacustrine sediments, and their bulk collapse into an inland rift-basin lake in the Moncton Subbasin, Canada, led to the entrainment of rare, almost complete, three-dimensionally preserved non-woody trees. Preservation of these unique fossils from the Albert Formation was a consequence of contemporaneous seismicity ..."

R.A. Gastaldo et al. (2024): Enigmatic fossil plants with three-dimensional, arborescent-growth architecture from the earliest Carboniferous of New Brunswick, Canada. Open access, Current Biology, 34: 1–12. https://doi.org/10.1016/j.cub.2024.01.011.
"... We present a new tree-crown architecture based on exceptional three-dimensional specimens
[...] this specimen shows that Early Carboniferous vegetation was more complex than realized, signaling that it was a time of experimental, possibly transitional and varied, growth architectures ..."
Note also:
3D-Fossilien skurriler Ur-Bäume entdeckt. (in German). By Tim Stonesifer, Bild der Wissenschaft.

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

R.A. Gastaldo et al. (2009): Ecological persistence in the Late Mississippian (Serpukhovian, Namurian A) megafloral record of the Upper Silesian Basin, Czech Republic. PDF file, Palaios, 24: 336-350.
See likewise 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.

! August von Gutbier (1835-1836):
Abdrücke und Versteinerungen des Zwickauer Schwarzkohlengebirges und seiner Umgebungen. Zwickau 1835-1836. In German. Provided by R. Daber, J. Meyer and P.B. Wendt, Botanischer Garten Berlin (Freie Universität Berlin). Excellent! Note the restored plates and
! photographs of Gutbier specimen.

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

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.

Illinois State Museum, Springfield:
Mazon Creek Plants.
Still available via Internet Archive Wayback Machine.

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.

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.

R.J. Knecht et al. (2024): Early Pennsylvanian Lagerstätte reveals a diverse ecosystem on a subhumid, alluvial fan. Open access, Nature Communications, 15. See likewise here.
"... We know relatively little about terrestrial ecosystems from the Early Pennsylvanian, which was a critical interval for the diversification of insects, arachnids, tetrapods, and seed plants
[...] Here we report a diverse Early Pennsylvanian trace and body fossil Lagerstätte (~320–318 Mya) from the Wamsutta Formation of eastern North America
[...] This site expands our knowledge of early terrestrial ecosystems and organismal interactions and provides ground truth for future phylogenetic reconstructions of key plant, arthropod, and vertebrate groups ..."

! 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.

M. Laaß and R. Rößler (2024): Kapitel 12: Tier-Pflanze-Interaktionen. PDF file, in German, in: R. Werneburg and J.W. Schneider (eds.): Die Rotliegend-Fauna des Thüringer Waldes.
See also Palökologie im Karbon und Perm des Thüringer-Wald-Beckens.

! 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.

J. Lies and R. Rößler (2024): Der Hornstein von Priefel - Ein Fossilvorkommen aus dem Perm bei Altenburg. PDF file, in German. Veröffentlichungen Museum für Naturkunde Chemnitz, 47: 15-58.
Note figure 4: Historic reconstruction of the Carboniferous vegetation (by Joseph Kuwassegs, 1850).

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.

! S.G. Lucas et al. (2023): An introduction to ice ages, climate dynamics and biotic events: the Late Pennsylvanian world. Open access, Geological Society, London, Special Publications, 535.
Note figure 2: Late Pennsylvanian palaeogeographical map.
Figure 5: Reconstructions of Desmoinesian and Missourian age peat-forming swamp vegetation.

! 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, Spektrum.de, 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).

A. Molina-Solís et al. (2024): Macrofloral biostratigraphy reflects late Carboniferous vegetation dynamics in the Nord-Pas-de-Calais Coalfield, France. Open access, Papers in Palaeontology, 10.

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.

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.

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. Free access, Review of Palaeobotany and Palynology, 308.
"... The resultant 67 photomicrographs were photo-stitched into one large image
[...] Single microspores and pollen within the ovule were photographed under oil at ×100 using multiple images at different focus depths and then Z-stacked ..."

! 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).

S. Schachat et al. (2023): Vegetational change during the Middle–Late Pennsylvanian transition in western Pangaea. Abstract, Geological Society, London, Special Publications, 535: 337-359.
"... Results indicate no substantive taxonomic turnover across the boundary. This stands in marked contrast to patterns in mid-Pangaean coal basins where there is a large wetland vegetational turnover.
[...] immediately following the boundary in New Mexico, and for approximately half of the Missourian Stage, floras previously dominated by hygromorphs become overwhelmingly dominated by mesomorphic/xeromorphic taxa ..."

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.

A.C. Scott (2024): Carboniferous wildfire revisited: Wildfire, post-fire erosion and deposition in a Mississippian crater lake. In PDF, Proceedings of the Geologists' Association, 135: 416-437.
See likewiswe here.
Note figure 6c–e. Scanning electron micrographs of charred Metaclepsydropsis.
! Figure 13f-h: Scanning electron micrograph of charcoalified pteridosperm leaf dissolved from Pettycur Limestone block.

! A.C. Scott (2024): Thirty Years of Progress in Our Understanding of the Nature and Influence of Fire in Carboniferous Ecosystems. In PDF, Fire, 7. 248. https://doi.org/10.3390/fire7070248.
See here as well.
Note figure 7: The interpretation of the Viséan East Kirkton environment highlighting the role of wildfire.
"... One of the basic problems was the fact that charcoal-like wood fragments, so often found in sedimentary rocks and in coals, were termed fusain and, in addition, many researchers could not envision wildfires in peat-forming systems. The advent of Scanning Electron Microscopy and studies on modern charcoals and fossil fusains demonstrated beyond doubt that wildfire residues may be recognized in rocks dating back to at least 350 million years ..."

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 (Maineet- Loire), South of the Armorican Massif, France. Open access, Botany Letters, 170: 183-193. DOI: 10.1080/23818107.2023.2183899.

! 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. (2023): First report of silicified wood from a late Pennsylvanian intramontane basin in the Pyrenees: systematic affinities and palaeoecological implications. Free access, Papers in Palaeontology, 9. doi: 10.1002/spp2.1524.
"... The specimens correspond to two types of arborescent plants, a calamitacean Equisetales (Arthropitys sp.) and a Cordaitales (Dadoxylon sp.). They provide information not available from the adpression flora found in this locality, such as growth patterns, interactions with fungi, and the presence of tyloses ..."

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.

M.L. Trivett and G.W. Rothwell (1991): Diversity among Paleozoic Cordaitales. In PDF, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 183: 289-305.

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.

X. Wang et al. (2024): New Discovery of Calamitaceae from the Cisuralian in Northwest China: Morphological Evolution of Strobilus. Open access, Biology, 13. https://doi.org/10.3390/biology13050347.
Note figure 6: The age distribution and evolution of calamitean strobili.
"... This paper is based on the detailed description of the macro- and microstructures of Calamites and Macrostachya ..."

J.P. Wilson et al. (2023): Physiological selectivity and plant–environment feedbacks during Middle and Late Pennsylvanian plant community transitions. Open access, Geological Society, London, Special Publications, 535: 361-382.
"... We find that three Pennsylvanian plant lineages – the medullosans, arborescent lycopsids and Sphenophyllum – contain high hydraulic conductivity but are vulnerable to drought-induced damage, whereas others are resistant, including stem group tree ferns and coniferophytes ..."

! 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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