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Permian Palaeobotany


A. Abu Hamad et al. (2017): Dicroidium bandelii sp. nov. (corystospermalean foliage) from the Permian of Jordan. In PDF, PalZ, 91: 641–648. See also here.

D. Agnihotri et al. (2024): Satpuraphyllum furcatum—a new genus and species of Peltaspermales foliage from the mid-Permian Barakar Formation of India. Free access, Alcheringa: An Australasian Journal of Palaeontology, DOI: 10.1080/03115518.2024.2415097.

! 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.

M. Backer et al. (2019): Frond morphology and epidermal anatomy of Compsopteris wongii (T. Halle) Zalessky from the Permian of Shanxi, China. Open access, PalZ.

! M. Barthel (2016): Die Rotliegendflora der Döhlen-Formation. PDF file, in German. Geologica Saxonica, 61: 105-238.

M. Barthel et al. (2010): Die Rotliegendflora des Weißig-Beckens. PDF file, in German. Geologica Saxonica, 56: 159-192.

Geologica Saxonica. Journal of Central European Geology. Senckenberg Naturhistorische Sammlungen Dresden, Abteilung Museum für Mineralogie und Geologie.

K. Bauer et al. (2014): Ginkgophytes from the upper Permian of the Bletterbach gorge (northern Italy). In PDF, see also here.

K. Bauer et al. (2013): The ginkgophytes from the German Kupferschiefer (Permian), with considerations on the taxonomic history and use of Baiera and Sphenobaiera. In PDF, Bulletin of Geosciences, 88: 539-556.

P. Blomenkemper et al. (2021): Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group. In PDF, Front. Earth Sci., 9: 652699. doi: 10.3389/feart.2021.652699.
See also here.

P. Blomenkemper et al. (2019): Cryptokerpia sarlaccophora gen. et sp. nov., an enigmatic plant fossil from the Late Permian Umm Irna Formation of Jordan. In PDF, PalZ, 93: 479–485. See also here.

P. Blomenkemper et al. (2018): A hidden cradle of plant evolution in Permian tropical lowlands. Abstract, Science, 362: 1414-1416. See also here (researchers from the University of Münster report on their findings), and there (Scinexx article, in German).

R. Chatterjee et al. (2014): Dwarfism and Lilliput effect: a study on the Glossopteris from the late Permian and early Triassic of India. In PDF, Current Science. See also here and there (abstract).

B.-Y. Chen et al. (2022): Anatomy of Stigmaria asiatica Jongmans et Gothan from the Asselian (lowermost Permian) of Wuda Coalfield, Inner Mongolia, North China. In PDF, Palaeoworld, 31: 311–323.
See also here.

I.C. Christiano De Souza et al. (2012): Permian bryophytes of Western Gondwanaland from the Paraná Basin in Brazil. In PDF, Palaeontology, 55: 229-241.

D. Chu et al. (2020): Ecological disturbance in tropical peatlands prior to marine Permian-Triassic mass extinction. Open access, Geology, 48: 288–292.

D. Chu et al. (2016): Biostratigraphic correlation and mass extinction during the Permian-Triassic transition in terrestrial-marine siliciclastic settings of South China. In PDF, Global and Planetary Change, 146: 67–88.
See also here.

! 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 & 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.

! J.A. Clement-Westerhof (1984): Aspects of Permian palaeobotany and palynology. IV. The conifer Ortiseia florin from the val gardena formation of the dolomites and the Vicentinian alps (Italy) with special reference to a revised concept of the Walchiaceae (Göppert) Schimper. In PDF, Review of Palaeobotany and Palynology, 41: 51-166. See also here.

M.P. D'Antonio et al. (2021): Primary tissues dominated ground-level trunk diameter in Sigillaria: evidence from the Wuda Tuff, Inner Mongolia. In PDF, Journal of the Geological Society. See also here.
Note figs. 1-4: in situ stump casts of Sigillaria from the earliest Permian.

A.-L. Decombeix et al. (2016): Bark anatomy of Late Permian glossopterid trees from Antarctica. Abstract, IAWA Journal, 37: 444-458. See also here (in PDF).

A.L. Decombeix (2010): Understanding the biology of high-latitude trees in a greenhouse world. In PDF, Palaios, 25: 423–425.
See also here.
Note figure 1C: Cross section of two roots (Vertebraria) showing growth rings and the typical air spaces in the wood.

C.G. Diedrich (2009): A coelacanthid-rich site at Hasbergen (NW Germany): taphonomy and palaeoenvironment of a first systematic excavation in the Kupferschiefer (Upper Permian, Lopingian). In PDF, Palaeobio. Palaeoenv., 89: 67-94.
Mapped taphonomy of plants (hinterland flora), invertebrates and fish vertebrates at six different planal levels on a 12 m2 area.

W.A. DiMichele et al. (2023): A paleontological perspective on ecosystem assembly rules in the terrestrial Paleozoic. Free access, Evolving Earth.
Note figure 1: Early Devonian (Emsian) flora from Gaspé, Canada.
Figure 2C: Edaphosaurus feeding on Supaia plants on stream bank, with background vegetation dominated by conifers. Early Permian (Wolfcampian/Asselian), New Mexico.

W.A. DiMichele et al. (2023): Two Early Permian Fossil Floras from the Arroyo de Alamillo Formation of the Yeso Group, Socorro County, New Mexico. In PDF, New Mexico Museum of Natural History and Science Bulletin, 94.
! Note figure 3d: Mud draped surface from L8741. Surface is covered by cavities interpreted either as raindrop imprints or gas-escape structures.
! Figure 6: Branch fragments preserved in a mud drape.

W.A. DiMichele et al. (2015): Early Permian fossil floras from the red beds of Prehistoric Trackways National Monument, southern New Mexico. In PDF, New Mexico Museum of Natural History and Science, Bulletin, 65: 129-139. See also here.
! Note fig. 3 and 4: Large mats of Walchia branches encased in claystones.

W.A. DiMichele et al. (2015): A compositionally unique voltzian conifer-callipterid flora from a carbonate-filled channel, lower Permian, Robledo Mountains, New Mexico, and its broader significance (Google books). In: S.G. Lucas & W.A. DiMichele (Eds.), Carboniferous-Permian transition in the Robledo Mountains, sounthern New Mexico. New Mexico Museum of National History and Sciences Bulletin (Vol. 65, pp. 123–128). See also here (PDF file).

W.A. DiMichele et al. (2013): Growth habit of the late Paleozoic rhizomorphic tree-lycopsid family Diaphorodendraceae: Phylogenetic, evolutionary, and paleoecological significance. Open access, American Journal of Botany, 100: 1-22.

! William A. DiMichele et al. (2008): The so-called "Paleophytic–Mesophytic" transition in equatorial Pangea. Multiple biomes and vegetational tracking of climate change through geological time. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 268: 152-163.
See likewise here (abstract).

W.A. DiMichele et al. (2008): The so-called "Paleophytic–Mesophytic" transition in equatorial Pangea. Multiple biomes and vegetational tracking of climate change through geological time. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 268: 152-163.
See likewise here (abstract), and there (still available via Internet Archive Wayback Machine).
! "... the evidence for a global “Paleophytic” vs. “Mesophytic” “vegetation” is simply unsubstantiated by the fossil record.
[...] The vegetational changes occurring in the late Paleozoic thus can be understood best when examined as spatial–temporal changes in biome-scale species pools responding to major global climate changes, locally and regionally manifested. ..."

W.A. DiMichele et al. (2007): A low diversity, seasonal tropical landscape dominated by conifers and peltasperms: Early Permian Abo Formation, New Mexico. In PDF, Review of Palaeobotany and Palynology, 145: 249-273.

! 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. (2005): Equisetites from the Early Permian of North-Central Texas. PDF file. In: Lucas, S.G. and Zeigler, K.E., (eds.), The Nonmarine Permian, New Mexico Museum of Natural Histojy and Science Bulletin, 30.
See also here.

! W.A. DiMichele et al. (2004): An unusual Middle Permian flora from the Blaine Formation (Pease River Group: Leonardian-Guadalupian Series) of King County, West Texas. In PDF, J. Paleont., 78: 765-782.
See also here.
Paper awarded with the Winfried and Renate Remy Award 2005 (Paleobotanical Section), 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): An Early Permian flora with Late Permian and Mesozoic affinities from north-central Texas. In PDF, Journal of Paleontology, 75: 449-460.
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 (1999): EVOLUTIONARY AND PALEOECOLOGICAL IMPLICATIONS OF TERRESTRIAL FLORAL CHANGES IN THE LATE PALEOZOIC TROPICS. Abstract, 1999 GSA Annual Meeting, Denver, Colorado; The Geological Society of America (GSA).
This expired link is now available through the Internet Archive´s Wayback Machine.

H.J. Falcon-Lang et al. (2015): Early Permian (Asselian) vegetation from a seasonally dry coast in western equatorial Pangea: Paleoecology and evolutionary significance. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 433: 158–173.

Z. Feng et al. (2024): Fossil evidence for silica biomineralisation in Permian lycophytes. Open access, National Science Review,

Z. Feng et al. (2022): Nurse logs: A common seedling strategy in the Permian Cathaysian Flora. In PDF, iScience, 25.
See also here.
"... We report seven coniferous nurse logs from lowermost to uppermost Permian strata of northern China that have been colonized by conifer and sphenophyllalean roots. These roots are associated with two types of arthropod coprolites and fungal remains. ..."

Z. Feng et al. (2020): From rainforest to herbland: New insights into land plant responses to the end-Permian mass extinction. Free access, Earth-Science Reviews.
Note fig. 8: Tomiostrobus sinensis Feng, whole plant reconstruction.
Note fig. 9: Reconstructions of the late Permian and Early Triassic vegetation in Southwest China.

Z. Feng et al.(2017): Leaf anatomy of a late Palaeozoic cycad. Biol. Lett., 13.

! Z. Feng et al. (2012): When horsetails became giants. Free access, Chinese Science Bulletin, 57: pages 2285–2288.
Reconstruction of the horsetail tree Arthropitys bistriata.

C.R. Fielding et al. (2022): Environmental change in the late Permian of Queensland, NE Australia: The warmup to the end-Permian Extinction. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 594.
See also here.
"... the time interval 257–252 Ma represented by the studied succession does not record a simple monotonic change in palaeoenvironmental conditions, but rather a series of intermittent stepwise changes towards warmer, and more environmentally stressed conditions leading up to the EPE [End-Permian Extinction] in eastern Australia. ..."

M.A. Flores-Barragan and M.P. Velasco-de Leon (2021): New records of Bjuvia and Nilssonia from the Permian of Mexico. In PDF, Palaeontologia Electronica, 24. See also here.

F. Fluteau et al. (2001): The Late Permian climate. What can be inferred from climate modelling concerning Pangea scenarios and Hercynian range altitude? PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 167: 39-71.
See also here.

! G. Forte et al. (2017): Conifer diversity in the Kungurian of Europe — Evidence from dwarf-shoot morphology. Abstract, Rev. Palaeobot. Palynol. See also here (in PDF).

R.A. Gastaldo and M.K. Bamford (2023): The influence of taphonomy and time on the paleobotanical record of the Permian–Trisssic transition of the Karoo basin (and elsewhere). In PDF, Journal of African Earth Sciences, 204.
See also here.

R.A. Gastaldo et al. (2017): Paleontology of the Blaauwater 67 and 65 Farms, South Africa: testing the Daptocephalus/Lystrosaurus biozone boundary in a stratigraphic framework. In PDF, Palaios, 34: 369–366. See also here (abstract).
"Contrary to the proposal that the Karoo Basin experienced a vegetational die off in the upper Daptocephalus biozone that was responsible for a phased extinction of vertebrates, our collections indicate that glossopterids and sphenophytes continued to colonize landscapes of the Lystrosaurus AZ".

! R.A. Gastaldo et al. (1996): Out of the Icehouse into the Greenhouse: A Late Paleozoic Analog for Modern Global Vegetational Change. In PDF, GSA Today 10: 1–7.
Note figure 1: Reconstruction of middle late Carboniferous tropical coal swamp.
Figure 2: Relation between global glaciation and vegetative change during the late Paleozoic in different tropical environments and the north and south temperate belts.
"... Patterns in the late Paleozoic provide us with one certainty: global warming presents plants with conditions that are markedly different from those found during periods of icehouse climate. The waxing and waning of glaciers are, in and of themselves, a climate-mode to which vegetations become attuned ..."

A.V. Gomankov (2022): Cycads in the Permian of the Subangara Region. In PDF, Paleontological Journal, 56: 317–326.
See likewise here.
Note figure 2: General form of cladosperms of Dioonitocarpidium.
Figure 5: Scheme of presumed evolutionary development of cladosperms in the cycads.
"... The significance of Dioonitocarpidium as a possible initial form in the evolution of the Cycadales in the Mesozoic and Cenozoic is discussed ..."

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.

E.L. Gulbranson et al. (2014): Leaf habit of Late Permian Glossopteris trees from high-palaeolatitude forests. In PDF, Journal of the Geological Society, London, 171: 493–507.
Note fig. 1: Comparison of modern climate and biomes with those reconstructed for the latest Permian climate and biomes.

E.L. Gulbranson et al. (2012): Permian polar forests: deciduousness and environmental variation. In PDF, Geobiology, 10: 479-495.
See also here.
Note upright permineralized stumps in figure 3 and 6.

! 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. Hamad et al. (2008): A Late Permian flora with Dicroidium from the Dead Sea region, Jordan. In PDF, Review of Palaeobotany & Palynology 149: 85-130.

C.J. Harper et al. (2018): Fungal sporulation in a Permian plant fragment from Antarctica. In PDF, Bulletin of Geosciences, 93: 13–26. Czech Geological Survey, Prague.

Xiaoyuan He et al. (2010): Anatomically Preserved Marattialean Plants from the Upper Permian of Southwestern China: The Trunk of Psaronius laowujiensis sp. nov. PDF file, Int. J. Plant Sci.. 171: 662-678.

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

D.E. Horton et al. (2010): Influence of high-latitude vegetation feedbacks on late Palaeozoic glacial cycles. In PDF, Nature Geoscience, 3, pages 572–577. See also here.
"... Glaciation during the late Palaeozoic era (340–250 Myr ago) is thought to have been episodic, with multiple, often regional, ice-age intervals, each lasting less than 10 million years.
... [We] suggest that vegetation feedbacks driven by orbital insolation variations are a crucial element of glacial–interglacial cyclicity.

V.S. Isaev et al. (2018): The fossil Permian plants from the Vorkuta series, Pechora Coal basin. Recent acquisitions in the collection of the Earth Science Museum at Lomonosov Moscow University. Moscow University Bulletin. Series 4. Geology. See also here (in PDF).
Note fig. 3: A giant Permian dragonfly produces the ovipositions on the shoot of a large equisetophyte.
Note Photo series 2, fig: 3: Paracalamites aff. frigidus Neuburg; two shoots preserved vertically within the layer, in situ.

A. Jasper et al. (2013): The burning of Gondwana: Permian fires on the southern continent - a palaeobotanical approach. In PDF, Gondwana Research, 24: 148-160.
See also here.

E.V. Karasev et al. (2018): The Late Permian (Lopingian) and Early Triassic flora of the Moscow Syneclise. Advances in Devonian, Carboniferous and Permian Research: Stratigraphy, Environments, Climate and Resources. Bologna, p. 144-154.

! H. Kerp et al. (2021, start on PDF-page 141): The fossil flora of the Dead Sea region, Jordan–A late Permian Garden of Delights. Journal of Palaeosciences, 70: 135–158.
Still available via Internet Archive Wayback Machine.

! 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 et al. (2006): Typical Triassic Gondwanan floral elements in the Upper Permian of the paleotropics. Geology, 34: 265-268. See also here (in PDF).

A.V. Khramov et al. (2023): The earliest pollen-loaded insects from the Lower Permian of Russia. In PDF, Biol. Lett., 19: 20220523.
See also here.
Note figure 2k: Artistic reconstruction of female Tillyardembia feeding on Pechorostrobus pollen organ (Rufloriaceae).

V.A. Krassilov and E.V. Karasev (2009): Paleofloristic evidence of climate change near and beyond the Permian-Triassic boundary. PDF file, Palaeogeogr. Palaeoclimatol. Palaeoecol., 284: 326-336.

! 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.

E. Kustatscher et al. (2024): A Kungurian flora from the Southern Alps (Northern Italy) yielding cuticles. Free access, Review of Palaeobotany and Palynology, 323.

E. Kustatscher et al. (2019): Did the Czekanowskiales already exist in the late Permian? Free access, PalZ.

E. Kustatscher et al. (2017): The Lopingian (late Permian) flora from the Bletterbach Gorge in the Dolomites, Northern Italy: a review. In PDF, Geo.Alp, 14.

Sunia Lausberg (2002): Neue Kenntnisse zur saarpfälzischen Rotliegendflora ... Abstract, PDF file, Thesis, Section of Palaeobotany in Muenster, Germany (in German). Go to: Kapitel III: Die Coniferen des Jungpaläozoikums..
Kapitel IV: Eine Coniferen-dominierte Flora aus dem Unterrotliegend von Alsenz, Saar-Nahe-Becken. See also here.

S. Lausberg and H. Kerp (2000): Eine Coniferen-dominierte Flora aus dem Unterrotliegend von Alsenz, Saar-Nahe-Becken, Deutschland. In PDF, Feddes Repertorium.

D. Li et al. (2022): Leaf scar and petiole anatomy reveal Pecopteris lativenosa Halle is a marattialean fern. In PDF, Geobios, 72–73: 37-53.
See also here.
"... reveal that Pecopteris lativenosa possesses Caulopteris-type stem, stewartiopterid petioles and rachises, and belongs to the Paleozoic Marattiales family Psaroniaceae. ..."

L. Liu et al. (2020): A whole calamitacean plant Palaeostachya guanglongii from the Asselian (Permian) Taiyuan Formation in the Wuda Coalfield, Inner Mongolia, China. Abstract, Review of Palaeobotany and Palynology. See also here (in PDF).
Please note the whole plant reconstruction in figure 18.

! C.V. Looy and I.A.P. Duijnstee (2019): Voltzian Conifers of the South Ash Pasture Flora (Guadalupian, Texas): Johniphyllum multinerve gen. et sp. nov., Pseudovoltzia sapflorensis sp. nov., and Wantus acaulis gen. et sp. nov. Abstract, International Journal of Plant Sciences, 181. See also here (in PDF).

C.V. Looy et al. (2016): Biological and physical evidence for extreme seasonality in central Permian Pangea. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 451: 210–226. See also here (in PDF).

! C.V. Looy et al. (2014): The late Paleozoic ecological-evolutionary laboratory, a land-plant fossil record perspective. In PDF, The Sedimentary Record, 12: 4-18. See also here.

S.G. Lucas et al. (2023): Paleoecological Significance of Trace Fossils and Fossil Plants from a New Locality in the Upper Part of the Abo Formation (Lower Permian), Socorro County, New Mexico. In PDF, New Mexico Museum of Natural History and Science Bulletin, 94.

S.G. Lucas (2023): Permophiles Perspective: Nonmarine Permian Biostratigraphy, Biochronology and Correlation. In PDF, Permophiles.
Note figure 1: Map of Pangea at 270 Ma.

! L. Luthardt et al. (2023): Cycadodendron galtieri gen. nov. et sp. nov.: An Early Permian Gymnosperm Stem with Cycadalean Affinity. Free access, International Journal of Plant Sciences, 184.
Note figure 10: Details of cycad-specific stem-anatomical features.
"... Cycadodendron galtieri gen. nov. et sp. nov. represents a petrified cycad stem of early Permian age providing the oldest-known evidence of cycad anatomy.
[...] The broad anatomical similarities of C. galtieri with other fossil and extant cycads demonstrate the early evolution of various cycad-specific anatomical features in the lower Permian ..."

! L. Luthardt et al. (2022): Upside-down in volcanic ash: crown reconstruction of the early Permian seed fern Medullosa stellata with attached foliated fronds. Open access, PeerJ, 10: e13051.
"... The upper part of a Medullosa stellata var. typica individual broke at its top resulting from the overload of volcanic ash and was buried upside-down in the basal pyroclastics. The tree crown consists of the anatomically preserved apical stem, ten attached Alethopteris schneideri foliated fronds with Myeloxylon-type petioles and rachises. ..."

L. Luthardt et al. (2018): Severe growth disturbances in an early Permian calamitalean – traces of a lightning strike? In PDF, Palaeontographica Abteilung B, 298: 1-22.
See also here.
! "... The special injury of the calamitalean described herein [...] exhibits an elongated to triangular shape, a central furrow, a scar-associated event ring of collapsed to distorted tracheids, and was ultimately overgrown by callus parenchyma. We suggest that this scar most likely was caused by a lightning strike ..."

! 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).

L. Luthardt and R. Rößler (2017): Fossil forest reveals sunspot activity in the early Permian. Abstract, Geology. See also here (in PDF).

L. Luthardt et al. (2016): Palaeoclimatic and site-specific conditions in the early Permian fossil forest of Chemnitz—Sedimentological, geochemical and palaeobotanical evidence. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 441: 627–652.
See also here.

C. Mays and S. McLoughlin (2019): Caught between mass extinctions - the rise and fall of Dicroidium. In PDF.

! S. McLoughlin et al. (2024): Evidence for saprotrophic digestion of glossopterid pollen from Permian silicified peats of Antarctica. Free access, Grana. https://doi.org/10.1080/00173134.2024.2312610.
"... we describe translucent bodies referable either to fungi (Chytridiomycota) or water moulds (Oomycetes) within the pollen of glossopterid gymnosperms and cordaitaleans, and fern spores from silicified Permian (Guadalupian–Lopingian) peats
[...] Our study reveals that the extensive recapture of spore/pollen-derived nutrients via saprotrophic digestion was already at play in the high-latitude ecosystems of the late Palaeozoic ..."

S. McLoughlin (2022): Late Permian flora of the Little River Coal Measures, northeastern Australia. In PDF, Geophytology, 50: 37–48.

! S. McLoughlin and R. Prevec (2021): The reproductive biology of glossopterid gymnosperms—A review. Free access, Review of Palaeobotany and Palynology, 295. See also here (in PDF).
! Note fig. 2: Diagramatic reconstructions of glossopterid pollen-bearing organs.

S. McLoughlin et al. (2018): Pachytestopsis tayloriorum gen. et sp. nov., an anatomically preserved glossopterid seed from the Lopingian of Queensland, Australia. Chapter 9, in PDF, in: M. Krings, C.J. Harper, N.R. Cuneo and G.W. Rothwell (eds.): Transformative Paleobotany Papers to Commemorate the Life and Legacy of Thomas N. Taylor.

S. McLoughlin (2017): Antarctica’s Glossopteris forests. In PDF, In: 52 More Things You Should Know About Palaeontology, eds. A. Cullum, A.W. Martinius. Nova Scotia: Agile Libre, p. 22-23. See also here.

S. McLoughlin et al. (2015): Paurodendron stellatum: A new Permian permineralized herbaceous lycopsid from the Prince Charles Mountains, Antarctica. In PDF, Review of Palaeobotany and Palynology, 220: 1-15. Reconstruction on PDF page 11.
See also here.

S. McLoughlin (2011): Glossopteris - insights into the architecture and relationships of an iconic Permian Gondwanan plant. In PDF, J. Botan. Soc. Bengal 65: 1-14.

! M.F. Miller et al. (2016): Highly productive polar forests from the Permian of Antarctica. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 441: 292–304. See also here (in PDF).

! I.P. Montañez et al. (2016): Climate, pCO2 and terrestrial carbon cycle linkages during late Palaeozoic glacial–interglacial cycles. In PDF, Nature Geoscience, 9: 824–828.
See also here.
Note figure 2: Consensus pCO2 curves defined by LOESS analysis of combined pedogenic carbonate- and fossil plant-based CO2 estimates.

! S.V. Naugolnykh (2016): Flora Permica. Plant World of Permian Period. Cis-Urals. PDF file, in Russian. Transactions of the Geological Institute, 612. Geological Institute of the Russian Academy of Sciences, Moscow.
The monograph deals with the Permian flora of Western Angaraland (Middle and South Cis-Urals). With many line drawings and plant reconstructions.
You may navigate through the paper starting from the table of contents, PDF page 334.

S.V. Naugolnykh (2014): Fossil Flora from the Aleksandrovskoe Locality (Lower Permian, Kungurian; Krasnoufimsk District of the Sverdlovsk Region): Taxonomical Composition, Taphonomy, and a New Lycopsid Representative. In PDF, Paleontological Journal, 48: 209–217. See also here (abstract).

S.V. Naugolnykh (2013): Permian ferns of western Angaraland. In PDF, Paleontological Journal, 47: 1379–1462.
See likewise here.

! S.V. Naugolnykh (2009): A new fertile Neocalamites from the Upper Permian of Russia and equisetophyte evolution. In PDF. Geobios, 42: 513-523. See also here.
Note fig. 5: Neocalamites tubulatus nov. sp.; reconstruction of the stems with the lateral strobilus in attachment (left) and lateral shoot scar in the node (right).

! 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 et al. (2022): Diversity and Stratigraphic Distribution of Sphenophytes in the Permian of the Paraná Basin, Brazil. In PDF, In: Iannuzzi, R., Rößler, R., Kunzmann, L. (eds.): Brazilian Paleofloras. Springer.
See also here.

R. Neregato et al. (2017): New petrified calamitaleans from the Permian of the Parnaíba Basin, central-north Brazil, part II, and phytogeographic implications for late Paleozoic floras. In PDF, Review of Palaeobotany and Palynology, 237: 37–61. See also here.
Note fig. 2 (on PDF page 16): The proposed reconstruction of Arthropitys tocantinensis sp. nov., drawn by F. Spindler, Freiberg).

R Neregato et al. (2015): New petrified calamitaleans from the Permian of the Parnaíba Basin, central-north Brazil. Part I. In PDF, Review of Palaeobotany and Palynology, 215: 23-45. See also here.
Note fig. 3 (on PDF page 15): The proposed reconstruction of Arthropitys isoramis sp. nov., drawn by F. Spindler, Freiberg).

Paläontologische Gesellschaft:
Fossil of the Year 2023.
About Medullosa stellata and fronds of the type Alethopteris schneideri. More information from the website in German.

Permophiles.
Newsletter of the Subcommission on Permian Stratigraphy.

A.G. Ponomarenko (2006): Changes in terrestrial biota before the Permian-Triassic ecological crisis. Abstract.

R. Prevec et al. (2022): South African Lagerstätte reveals middle Permian Gondwanan lakeshore ecosystem in exquisite detail. Open access, Communications Biology, 5.
Note figure 1: Climatic zones for the Wordian of Pangea including locations of middle Permian fossil insect discoveries.
Figure 6: Reconstruction of a middle Permian lakeshore palaeoenvironment.

R. Prevec et al. (2009): Portrait of a Gondwanan ecosystem: A new late Permian fossil locality from KwaZulu-Natal, South Africa. Abstract, Review of Palaeobotany and Palynology, 156: 454-493. See also here, or there (PDF files).

! P. McAllister Rees (2002): Land-plant diversity and the end-Permian mass extinction. PDF file, Geology, 30: 827-830. See also here (abstract).

P.M.A. Rees et al. (1999): Permian climates: Evaluating model predictions using global paleobotanical data. In PDF, Geology, 27: 891-894. See also here.

P.M. Rees et al. (2002): Permian Phytogeographic Patterns and Climate Data/Model Comparisons. PDF file, Journal of Geology, 110, 1–31.
See also here.

G.J. Retallack and E.S. Krull (1999): Landscape ecological shift at the Permian-Triassic boundary in Antarctica. In PDF, Australian Journal of Earth Sciences.
Now provided by the Internet Archive´s Wayback Machine.

D. Rockenbach Boardman et al. (2016): A new genus of Sphenopsida from the Lower Permian of the Paraná Basin, Southern Brazil. In PDF, Review of Palaeobotany and Palynology, 233: 44–55. See also here and there.

R. Rößler (2019): Der Wald aus Stein unter Chemnitz – einzigartiges „Pompeji des Erdaltertums“. In German, PDF file. Kalenderblatt April 2019, Online-Plattform der Professur Geschichte Europas im Mittelalter und in der Frühen Neuzeit an der Technischen Universität Chemnitz.
See also here.

R. Rößler et al. (2015): Der Versteinerte Wald Chemnitz - Momentaufnahme eines vulkanisch konservierten Ökosystems aus dem Perm (Exkursion L am 11. April 2015). PDF file, in German. The petrified forest of Chemnitz - A snapshot of an early Permian ecosystem preserved by volcanism. Jber. Mitt. oberrhein. geol. Ver., N.F. 97.

R. Rößler (2014): Die Bewurzelung permischer Calamiten: Aussage eines Schlüsselfundes zur Existenz freistehender baumförmiger Schachtelhalmgewächse innerhalb der Paläofloren des äquatornahen Gondwana. PDF file, in German. The roots of Permian calamitaleans - a key find suggests the existence of free-stemmed arborescent sphenopsids among the low latitude palaeofloras of Gondwana. Freiberger Forschungshefte, C 548.

! R. Rößler et al. (2012): The largest calamite and its growth architecture - Arthropitys bistriata from the Early Permian Petrified Forest of Chemnitz. In PDF, Review of Palaeobotany and Palynology, 185: 64-78.
The link is to a version archived by the Internet Archive´s Wayback Machine.

R. Rößler and R. Noll (2006): Sphenopsids of the Permian (I): The largest known anatomically preserved calamite, an exceptional find from the petrified forest of Chemnitz, Germany. Abstract, Review of Palaeobotany and Palynology, 140: 145–162. See also here (in PDF).

Ronny Rößler & Robert Noll (website hosted by fossilien-journal.de): Calamitea COTTA 1832. Fossile Pflanze zwischen Historie und aktueller Forschung. PDF file, in German. Snapshot taken by the Internet Archive´s Wayback Machine.

R. Rößler, (2006): Einzigartig und dennoch ausgestorben - Die Schachtelhalm-Giganten des Perms (in German). In PDF, Fossilien, 23: 87-92. Provided by the Internet Archive´s Wayback Machine.

R. Rößler and M. Barthel(1998): Rotliegend taphocoenoses preservation favoured by rhyolitic explosive volcanism. In PDF, Freiberger Forschungshefte C, 474: 59–101. See also here.

K. Ruckwied et al. (2015): Palynological records of the Permian Ecca Group (South Africa): Utilizing climatic icehouse-greenhouse signals for cross basin correlations. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 413: 167-172.
The link is to a version archived by the Internet Archive´s Wayback Machine.
See also here.

P.E. Ryberg and E.L. Taylor, Department of Ecology and Evolutionary Biology; Natural History Museum and Biodiversity Research Center, University of Kansas, Lawrence: Silicified wood from the Permian and Triassic of Antarctica: Tree rings from polar paleolatitudes. PDF file, Geological Survey and The National Academies; USGS OF-2007-1047, Short Research Paper 080.

! Sächsische Landesamt für Umwelt und Geologie (2006): Das Döhlener Becken bei Dresden - Geologie und Bergbau. PDF file, in German. Bergbau in Sachsen, vol. 12. See especially PDF page 30: Makroflora und zugehörige "in situ"-Sporen (by M. Barthel).

A. Saxena et al. (2022): Early Permian macrofloral diversity in Indian Gondwana: Evidence from Talchir Formation of Singrauli coalfield, Son–Mahanadi valley basin, central India. In PDF, Journal of Earth System Science, 131.
See also here.

A.B. Schwendemann (2024): A leaf economics analysis of high-latitude Glossopteris leaves using a technique to estimate leaf mass per area. Evolving Earth, 2.
"... An analysis of the leaf mass per area (LMA) of late Permian Glossopteris leaves from Antarctica gives several insights into how these fossil leaves fit into functional groups and habitats compared to extant plants.
[...] When combined with the known effects of high CO2 and continuous light conditions on leaf LMA [leaf mass per area], the data suggest that the glossopterids living in these polar latitudes had seasonally deciduous leaves and adaptations that allowed them to thrive in a continuous light environment ..."

L.J. Seyfullah et al. (2010): Resolving the systematic and phylogenetic position of isolated ovules: a case study on a new genus from the Permian of China. In PDF, Botanical Journal of the Linnean Society, 164: 84–108. See also here.

! X. Shi (2016): Fossil plants and environmental changes during the Permian-Triassic transition in Northwest China. Doctoral dissertation, Université Pierre et Marie Curie, China University of Geosciences Wuhan. See also here (abstract).

X. Shi (2016): Fossil plants and environmental changes during the Permian-Triassic transition in Northwest China. Thesis, Université Pierre et Marie Curie,Paris VI. See also here.

! G.R. Shi and J.B. Waterhouse (2010): Late Palaeozoic global changes affecting high-latitude environments and biotas: an introduction. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 298: 1-16.

S.S.T. Simon et al. (2018): An exhumed fine-grained meandering channel in the lower Permian Clear Fork Formation, north-central Texas: Processes of mud accumulation and the role of vegetation in channel dynamics. In PDF, Int. Assoc. Sedimentol., Spec. Publ., 48: 149–172.
See also here.
"... weakly laminated mudstone with desiccation cracks contains leaves and seeds of Evolsonia texana, marattialean foliage and Taeniopteris sp., with root traces penetrating the leaves. ..."

B.J. Slater et al. (2015):

S.S.T. Simon et al. (2016): An abandoned-channel fill with exquisitely preserved plants in redbeds of the Clear Fork Formation, Texas, USA: an Early Permian water-dependent habitat on the arid plains of Pangea. In PDF, J. Sed. Res., 86, 944–964. See also here.
Note fig. 11: Goethite petrification of cellular structure of plant remains.

A high-latitude Gondwanan lagerstätte: The Permian permineralised peat biota of the Prince Charles Mountains, Antarctica. In PDF, Gondwana Research, 27: 1446-1473. See also here (abstract).

B.J. Slater et al. (2013): Peronosporomycetes (Oomycota) from a Middle Permian Permineralised Peat within the Bainmedart Coal Measures, Prince Charles Mountains, Antarctica.

J.M. Souza and R. Iannuzzi (2012): Dispersal Syndromes of fossil Seeds from the Lower Permian of Paraná Basin, Rio Grande do Sul, Brazil. Click: "PDF in English". An. Acad. Bras. Ciênc., 84: 3-68.

R. Spiekermann et al. (2018): A remarkable mass-assemblage of lycopsid remains from the Rio Bonito Formation, lower Permian of the Paraná Basin, Rio Grande do Sul, Brazil. In PDF, Palaeobiodiversity and Palaeoenvironments, 98: 369–384. See also here.

A.K. Srivastava and R. Srivastava (2016): Glossopteridales: An intricate group of plants. In PDF, The Palaeobotanist, 65: 159–167.

A.K. Srivastava and D. Agnihotri (2010): Dilemma of late Palaeozoic mixed floras in Gondwana. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology. See also here (abstract).

R. Tewari et al. (2017): The Glossopteris flora of Manuguru Area, Godavari Graben, Telangana, India. In PDF, Palaeobotanist, 66: 17–36.

! R. Tewari et al. (2015): Glossopteris flora in the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica: Implications for paleogeography, paleoclimatology, and biostratigraphic correlation. Abstract, GR Focus Review, Gondwana Research, 28: 905-932. See also here (in PDF).

D. Uhl and H. Kerp (2020): 4.2.3 Die terrestrische Makroflora des Zechsteins. PDF file, in German. Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften 89: 83-91. In: Deutsche Stratigraphische Kommission.
See also here.

D. Uhl (2013): The paleoflora of Frankenberg/Geismar (NW-Hesse, Germany) - a largely unexplored "treasure chest" of anatomically preserved plants from the Late Permian (Wuchiapingian) of the Euramerican floral province. PDF file; In: Lucas, S.G., et al. eds., The Carboniferous-Permian Transition. New Mexico Museum of Natural History and Science. Bulletin, 60, 433-443.

K. Unger Baillie (March 12, 2021): ‘Pompeii of prehistoric plants’ unlocks evolutionary secret. Penn Today.

I.M. Van Waveren et al. (2021): Climate-driven palaeofloral fluctuations on a volcanic slope from the low latitudes of the Palaeotethys (early Permian, West Sumatra). In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 579. See also here.

M. Wan et al. (2016): A typical Euramerican floral element from the Shanxi Formation (Cisuralian, lower Permian) in the Wuda Coal Field, Inner Mongolia, North China. Palaeobiodiversity and Palaeoenvironments, 96: 507–515.
Provided by the Internet Archive´s Wayback Machine.
See also here.

J. Wang et al. (2021): Ancient noeggerathialean reveals the seed plant sister group diversified alongside the primary seed plant radiation: Open access, Proceedings of the National Academy of Sciences, 118, e2013442118.
Note fig. 2: Reconstruction of the aerial parts of Paratingia wuhaia from the early Permian of China.

J. Wang et al. (2012): Permian vegetational Pompeii from Inner Mongolia and its implications for landscape paleoecology and paleobiogeography of Cathaysia. In PDf, PNAS, 109: 4927-4932. Reconstructions of peat-forming forests of earliest Permian age in fig. 4 and 5.

Jun Wang et al. (2012): Permian vegetational Pompeii from Inner Mongolia and its implications for landscape paleoecology and paleobiogeography of Cathaysia. In PDF, PNAS. See also: Ash-covered forest is "Permian Pompeii" (S. Perkins, Nature).
Penn researcher helps discover and characterize a 300-million-year-forest.
The Lost Forest.

Jun Wang and Hermann W. Pfefferkorn (2010): Nystroemiaceae, a new family of Permian gymnosperms from China with an unusual combination of features. PDF file, Proc. R. Soc., B, 277: 301-309. See also here.

S.-J. Wang et al. (2017): Anatomically preserved "strobili" and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Free access, Am. J. Bot., 104: 127-149.

Wang Ziqiang and Zhang Zhiping (1998): Gymnosperms on the eve of the terminal Permian mass extinction in North China and their survival strategies. In PDF, Chinese Science Bulletin, 43: 889-897.

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 ..."

M.M. Windell (2024): A Permian permineralised peat reveals high spatial and temporal variation in plant assemblage. In PDF. Degree Project in Physical Geography and Quaternary Geography, Stockholm University.
See here as well.
Note figure 10: Reconstruction of the rift-valley-bound mid-Permian forest swamp ecosystem of East Antarctica, at the beginning of autumn.

! Q. Wu et al. (2021): High-precision U-Pb age constraints on the Permian floral turnovers, paleoclimate change, and tectonics of the North China block. Free access, Geology. See also here.
"... The great loss of highly diverse and abundant Cathaysian floras and the widespread invasion of the Angaran floras under arid climate conditions in the North China block happened during the late Cisuralian to Guadalupian, but its exact timing is uncertain due to the long hiatus. ..."

W. Zhou et al. (2022): Diodonopteris virgulata sp. nov., a climbing fern from the early Permian Wuda Tuff Flora and its paleoecology. In PDF, Review of Palaeobotany and Palynology, 304.
See also here.

W.-M. Zhou et al. (2021): An upright psaroniaceous stump and two surrounding pecopteroids from the early Permian Wuda Tuff Flora. In PDF, Palaeoworld, 30: 451-460.
See also here.
Note figure 2: Morphology and measurements of the Psaronius stump.

W. Zhou et al. (2020): Yangopteris ascendens (Halle) gen. et comb. nov., a climbing alethopterid pteridosperm from the Asselian (earliest Permian) Wuda Tuff Flora. In PDF, Review of Palaeobotany and Palynology. See also here.
















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