Links for Palaeobotanists

Home / Preservation & Taphonomy / Cuticles

Taphonomy in General
Plant Fossil Preservation and Plant Taphonomy
Collecting Bias: Our Incomplete Picture of the Past Vegetation
Three-Dimensionally Preserved Plant Compression Fossils
Pith Cast and "in situ" Preservation
Bacterial Biofilms (Microbial Mats)
Permineralized Plants and the Process of Permineralization
Petrified Forests
Pyrite Preservation
Molecular Palaeobotany
Upland and Hinterland Floras
Abscission and Tissue Separation in Fossil and Extant Plants
Leaf Litter and Plant Debris
Log Jams and Driftwood Accumulations
Wound Response in Trees
Fungal Wood Decay: Evidence from the Fossil Record

! Transfer Technique@
! Stomatal Density@
! Chemotaxonomy and Chemometric Palaeobotany@
! Overviews of Plant Fossil Lagerstätten and Their Palaeoenvironments@
Plant Anatomy@
Introductions to both Fossil and Recent Plant Taxa@


A.M.B. Abu Hamad et al. (2019): The first record of Dicroidium from the Triassic palaeotropics based on dispersed cuticles from the Anisian Mukheiris Formation of Jordan. In PDF, PalZ, 93. 487–498. See also here.

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.

American Society of Plant Biologists, The Plant Cell Online: Leaf Development 1 and Leaf Development 2 (Cell proliferation and differentiation). Still available via Internet Archive. Lecture notes, PDF files. For PowerPoint Slide Presentations see here.

S. Archangelsky (1968): Studies on Triassic fossil plants from Argentina. IV. The leaf genus Dicroidium and its possible relation to Rhexoxylon stems. PDF file, Palaeontology.
The link is to a version archived by the Internet Archive´s Wayback Machine.

Anne-Marie Aucour et al. (2009): Insights into preservation of fossil plant cuticles using thermally assisted hydrolysis methylation. Abstract, Organic Geochemistry, 40: 784-794.

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.

G. Barale et al. (2005): A fossil peat deposit from the Late Triassic (Carnian) of Zimbabwe with preserved cuticle of Pteridospermopsida and Ginkgoales, and its geological setting G Barale. In PDF, Palaeont. afr., 41: 89-100.
See also here.

M. Barbacka et al. (2022): Early Jurassic coprolites: insights into palaeobotany and the feeding behaviour of dinosaurs. In PDF, Papers in Palaeontology.
See also here.

M. Barbacka (2009): Sphenophyta from the Early Jurassic of the Mecsek Mts., Hungary. Snapshot taken by the Internet Archive´s Wayback Machine. PDF file, Acta Palaeobotanica 49: 221-231. Cuticle showing stomata of Equisetites columnaris in fig. 15!

! M. Barbacka (1994): Komlopteris Barbacka, gen. nov., a segregate from Pachypteris Brongniart. In PDF, Review of Palaeobotany and Palynology, 83: 339-349.
See likewise here.

R.S. Barclay et al. (2013): New methods reveal oldest known fossil epiphyllous moss: Bryiidites utahensis gen. et sp. nov.(Bryidae). In PDF, American Journal of Botany, 100: 2450-2457.
Using the zstacking software package Automontage (Syncroscopy, Cambridge, UK).

! R. Barclay, P. Wilf, D. Dilcher, A. Sokoloff, J. Leon-Guerrero & C. Thurman: Cuticle Database. The Cuticle Database Project aims to promote the understanding and identification of living and fossil plants. This project is a collaborative effort between researchers at Northwestern University, The Field Museum, the Florida Museum of Natural History, and Pennsylvania State University. See also here:
! R. Barclay, et al. (2007): The cuticle database: developing an interactive tool for taxonomic and paleoenvironmental study of the fossil cuticle record. PDF file, In: Jarzen, D. M., Steven, R., Retallack, G. J. and Jarzen, S. A. (eds.), Advances in Angiosperm Paleobotany and Paleoclimatic Reconstruction, Contributions Honouring David L. Dilcher and Jack A. Wolfe, Courier Forschungsinstitut Senckenberg, Frankfurt, pgs. 39-56.

M.R. Barone Lumaga et al. (2015): Epidermal micromorphology in Dioon: did volcanism constrain Dioon evolution? In PDF, Botanical Journal of the Linnean Society, 2015, 179: 236–254. See also here.

Terese Barta, UW-Stevens Point, Mike Clayton, UW-Madison, Dave Hillier, UW-Stevens Point, Brad Mogen, UW-River Falls, Jim Perry, UW-Fox Valley, Jan Phelps, UW-Baraboo, Patricia Ploetz, UW-Stevens Point, Tom Volk, UW-La Crosse, & Bob Wise, UW-Oshkosh (page hosted by BioWeb, University of Wisconsin): UW-System Botanical Image Library. This site is devoted to making botanical images available and easily accessible for educational use. They aspire to provide a stable, maintained library of non-proprietary images that can be easily referenced by botanists designing their own educational applications. Go to: Leaf Anatomy, Modified Leaves (now via wayback).

A. Bartiromo (2012): The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions: A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants. Dissertation, Université Claude Bernard, Lyon.

M.E.P. Batista et al. (2017): New data on the stem and leaf anatomy of two conifers from the Lower Cretaceous of the Araripe Basin, northeastern Brazil, and their taxonomic and paleoecological implications. Open access, PLoS ONE, 12.

Ernst-Georg Beck, Zentrale für Unterrichtswesen: Biokurs 2001 (in German). Go to: Pflanzenanatomie und Photosynthese, Aufbau eines typischen Laubblattes.

! J.A. Berry et al. (2010): Stomata: key players in the earth system, past and present. Abstract, Current opinion in plant biology, 13: 232–239. See also here (in PDF).

C. Blanco-Moreno (2021): Preparation protocols for SEM visualization of charred fossil plants: the case of Weichselia reticulata pinnule anatomy. In PDF, Spanish Journal of Palaeontology, 36.

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).

P. Blomenkemper et al. (2016): Cuticular analysis of conifers from the Keuper (Triassic) of Franconia, southern Germany. Abstract, starting on PDF page 220.
Abstracts, XIV International Palynological Congress, X International Organisation of Palaeobotany Conference, Salvador, Brazil.

B. Bomfleur et al. (2018): Polar Regions of the Mesozoic-Paleogene Greenhouse World as Refugia for Relict Plant Groups. Chapter 24, 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.

! J.G. Bornemann (1856): Über organische Reste der Lettenkohlengruppe Thüringens. Ein Beitrag zur Fauna und Flora dieser Formation, besonders über fossile Cycadeen, nebst vergleichenden Untersuchungen über die Blattstruktur der jetztweltlichen Cycadeengattungen. In German, provided by Google books.
A pioneering depiction of dispersed fossil cuticles from the Triassic (Ladinian) of Germany.

Ilma Brewer, Robyn Overall, Nicholas Skelton, & Mark Curran, School of Biological Sciences The University of Sydney, Australia: The Revision Modules in Plant Anatomy. Snapshot taken by the Internet Archive´s Wayback Machine. A photomicrographic overview of the major plant tissues and organs, with glossary.

D.E.G. Briggs (1999): Molecular taphonomy of animal and plant cuticles: selective preservation and diagenesis. PDF file, Phil. Trans. R. Soc. Lond. B,354: 7-17. See also here.

T.J. Brodribb and S.A.M. McAdam (2017): Evolution of the stomatal regulation of plant water content. Open access, Plant Physiology, 175: 639–649.

! T.J. Brodribb et al. (2016): Xylem and stomata, coordinated through time and space. Abstract, Plant Cell and Environment, 40: 872–880. See also here (in PDF).

! M.J.M. Brown and G.J. Jordan (2023): No cell is an island: characterising the leaf epidermis using EPIDERMALMORPH, a new R package. Open access, New Phytologist, 237: 354–366.
"... we present a method to characterise individual cell size, shape (including the effect of neighbouring cells) and arrangement from light microscope images. We provide the first automated characterisation of cell arrangement ..."
Download the R package (quantifying and analysing epidermal cell shape, size and spatial arrangement),
and the manual.

M.A. Carizzo et al. (2019): Cuticle ultrastructure in Brachyphyllum garciarum sp. nov (Lower Cretaceous, Argentina) reveals its araucarian affinity. Abstract, Review of Palaeobotany and Palynology, 269: 104-128. See also here (in PDF).

Note fig. 7: Brachyphyllum garciarum sp. nov. Three-dimensional reconstruction of the cuticles.

! E.M. Carlisle et al. (2021): Experimental taphonomy of organelles and the fossil record of early eukaryote evolution. Open access, Science Advances, 7: eabe9487.
Note fig. 4A: Fossil of a Zelkova leaf from the Miocene Succor Creek Formation showing a chloroplast adpressed to the cell wall.

B. Chefetz (2007): Decomposition and sorption characterization of plant cuticles in soil. In PDF, Plant and Soil, 298: 21-30.

! J.W. Clark et al. (2022): The origin and evolution of stomata. Free access, Curr. Biol., 6: R539-R553. doi: 10.1016/j.cub.2022.04.040.
See likewise here.
! Note figure 1: The phylogenetic context for stomatal origins and evolution.
Figure 4: The diversity of stomatal responses among land plants.

Richard Cowen, Department of Geology, University of California, Davis, CA: History of Life, Third Edition.
Go to: Preservation and Bias in the Fossil Record.
These expired links are now available through the Internet Archive´s Wayback Machine.

Robert Roy Cowie, San Marcos, TX: Gas Exchange Characteristics of an Early Cretacerous Conifer, Pseudofrenelopsis varians, (Cheirolepidiaceae), and its inferred Paleoecology. (via webback machine). Go to: Preparation of Materials.

Richard Crang, University of Illinois at Urbana-Champaign, Andrey Vassilyev, St. Petersburg State University, Russia (McGraw-Hill Higher Education): Plant Anatomy. A website that supports the Electronic Plant Anatomy CD-ROM. An instructor view provides links to dynamic cartoons viewable using the Macromedia Flash Player. Go to: "Stomata" (opening and closing stomata), and "Leaf Structure".

Judith L. Croxdale, Department of Botany, University of Wisconsin, Madison (website hosted by Biology Online):
Stomatal patterning in angiosperms.
Still available through the Internet Archive´s Wayback Machine.
Stomatal pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views are discussed.

! E. Cullen and P.J. Rudall (2016): The remarkable stomata of horsetails (Equisetum): patterning, ultrastructure and development. Abstract, Annals of Botany, 118: 207–218.
See also here (in PDF).

John D. Curtis, Biology Department, University of Wisconsin; Nels R. Lersten, Department of Botany, Iowa State University, and Michael D. Nowak, Biology Department, University of Wisconsin: Photographic Atlas of Plant Anatomy. Go to: Epidermis and Stomates.

R.G. Daly and R.A. Gastaldo (2010): The effect of leaf orientation to sunlight on stomatal parameters of Quercus rubra around the Belgrade Lakes, central Maine. PDF file, Palaios, 25: 339-346.
See likewise here.

J.A. D´Angelo et al. (2012): Compression map, functional groups and fossilization: A chemometric approach (Pennsylvanian neuropteroid foliage, Canada). Abstract, International Journal of Coal Geology.

I. Degani-Schmidt and M. Guerra-Sommer (2019): Epidermal morphology of the cordaitalean leaf Noeggerathiopsis brasiliensis nom. nov. from the southern Paraná Basin (Lower Permian, Rio Bonito Formation) and paleoenvironmental considerations. In PDF, Braz. J. Geol., 49. See also here.

G.M. Del Fueyo et al. (2019): Permineralized conifer-like leaves from the Jurassic of Patagonia (Argentina) and its paleoenvironmental implications. Anais da Academia Brasileira de Ciências (Annals of the Brazilian Academy of Sciences), 91: (Suppl. 2): e20180363. See also

Elias De Leon and Brian Zhou, NewsWatch, National Geographic: In a High School Lab, Glimpses of an Ancient Climate.
This expired link is still available through the Internet Archive´s Wayback Machine.

! D.L. Dilcher (1974): Approaches to the identification of angiosperm leaf remains. In PDF, The Botanical Review, 40: 1–157. Also availabe via here (in PDF).
See also here.
"... Many techniques for the study of the morphology of modern and fossil leaves are included in this paper as well as tables outlining features of leaf venation and the epidermis ..."

! 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.

C. Dong et al. (2022): Leaves of Taxus with cuticle micromorphology fromthe Early Cretaceous of eastern Inner Mongolia, Northeast China. In PDF, Review of Palaeobotany and Palynology, 298.
See also here.

J.M. Drovandi et al. (2022): Dicroidium (Zuberia) zuberi (Szajnocha) Archangelsky from exceptional Carnian leaf litters of the Ischigualasto Formation, westernmost Gondwana. In PDF, Historical Biology, 34.
See also here.

M. Eberlein (2015): Bestimmungs- und Verbreitungsatlas der Tertiärflora Sachsens – Angiospermenblätter und Ginkgo. PDF file (in German). Thesis, University of Dresden (in German). First part of a reference book of the Tertiary flora of Saxony.
See also here.

! D. Edwards, H. Kerp and H. Hass (1998): Stomata in early land plants: an anatomical and ecophysiological approach. In PDF, Journal of Experimental Botany, Vol. 49, Special Issue, pp. 255–278.

C. Elliott-Kingston et al. (2014): Damage structures in leaf epidermis and cuticle as an indicator of elevated atmospheric sulphur dioxide in early Mesozoic floras. In PDF, Review of Palaeobotany and Palynology, 208: 25-42.

Beth Ellis et al. (2009): Manual of Leaf Architecture. Book announcement. The link is to a version archived by the Internet Archive´s Wayback Machine.
! See also here and there.

Encyclopedia of Earth. An electronic reference about the Earth, its natural environments, and their interaction with society. Go to: What are stomata? About stomatal density, size and shape, physiological function of stomata, optimal size of stomatal apertures, and stomatal conductance. More botany articles here, and there (all titles A-Z).

H. Failmezger et al. (2013): Semi-automated 3D Leaf Reconstruction and Analysis of Trichome Patterning from Light Microscopic Images. In PDF, see also here.

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

! V. Fernández et al. (2016): Cuticle Structure in Relation to Chemical Composition: Re-assessing the Prevailing Model. Open access, Front. Plant Sci., 31.

K.C. Fetter et al. (2018): StomataCounter: a deep learning method applied to automatic stomatal identification and counting. In PDF, bioRxiv. See also here

Ben Fletcher, Department of Animal and Plant Sciences, University of Sheffield:
The role of stomata in the early evolution of land plants.
How the atmosphere affects plants.
These expired links are available through the Internet Archive´s Wayback Machine.

Jennifer Forman, Department of Biology, University of Massachusetts, Boston: Land of the Glandular Trichomes. A microscopic look at plants in the Lamiaceae family.

Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine:
Navigate via: Notes for a Course in Paleobotany.
Websites outdated. Download versions archived by the Internet Archive´s Wayback Machine.

! R.A. Gastaldo and J.R. Staub (1999): A mechanism to explain the preservation of leaf litter lenses in coals derived from raised mires. PDF file, Palaeogeography Palaeoclimatology Palaeoecology, 149: 1-14. See also here.
! Note figure 5: Illustration of mechanism proposed for preservation of structurally preserved leaves in peat accumulations in raised mires throughout the Phanerozoic.

S. García Álvarez et al. (2009): The value of leaf cuticle characteristics in the identification and classification of Iberian Mediterranean members of the genus Pinus. In PDF, J. Linn. Soc., 161: 436–448.

J.J. Glas et al. (2012): Plant Glandular Trichomes as Targets for Breeding or Engineering of Resistance to Herbivores. In PDF, Int. J. Mol. Sci., 13: 17077-17103.

Govindjee, Department of Plant Biology , University of Illinois at Urbana-Champaign, Urbana:
The Photosynthesis Page Govindjee. Note especially:
! Teaching Material.

Bruce W. Grant and Itzick Vatnick, Teaching Issues and Experiments in Ecology (TIEE). This is a project of the Education and Human Resources Committee of the Ecological Society of America: Environmental Correlates of Leaf Stomata Density. The technique of making clear nail polish impressions of leaf stomata.

O.R. Green: Extraction Techniques for Palaeobotanical and Palynological Material. Abstract, pp 256-287. A Manual of Practical Laboratory and Field Techniques in Palaeobiology.

G. Guignard et al. (2024): TEM and EDS characterization in a Bennettitalean cuticle from the Lower Cretaceous Springhill Formation, Argentina. Free access, Review of Palaeobotany and Palynology, 320.
Note figure 7: Three-dimensional reconstruction of lower and upper cuticles of Ptilophyllum eminelidarum.
"New cuticle samples from the bennettitalean Ptilophyllum eminelidarum were herein studied using the combination of light microscopy (LM), scanning and transmission electron microscopy (SEM, TEM), and element analysis by Energy Dispersive Spectroscopy (EDS) ..."

G. Guignard (2021): Method for ultrastructural fine details of plant cuticles by transmission electron microscopy. In PDF, MethodsX, 8.
See also here.

! G. Guignard (2019): Thirty-three years (1986–2019) of fossil plant cuticle studies using transmission electron microscopy: A review. Abstract, Review of Palaeobotany and Palynology, 271. See also here (in PDF).

! N.S. Gupta et al. (2006): Reinvestigation of the occurrence of cutan in plants: implications for the leaf fossil record. Abstract and references, Paleobiology, 32: 432-449.

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

T.M. Harris: The Problems of Jurassic Palaeobotany. In PDF.

M. Haworth et al. (2023): The functional significance of the stomatal size to density relationship: Interaction with atmospheric [CO2] and role in plant physiological behaviour. Open access, Science of The Total Environment, 863.
"... Angiosperms generally possessed higher densities of smaller stomata that corresponded to a greater degree of physiological stomatal control consistent with selective pressures induced by declining [CO2] over the past 90 Myr. Atmospheric [CO2] has likely shaped stomatal size and density relationships alongside the interaction with stomatal physiological behaviour ..."

M. Haworth and A. Raschi (2014): An assessment of the use of epidermal micro-morphological features to estimate leaf economics of Late Triassic-Early Jurassic fossil Ginkgoales. In PDF, Review of Palaeobotany and Palynology, 205: 1-8.

M. Haworth et al. (2011): Stomatal control as a driver of plant evolution. In PDF, J. Exp. Bot., 62: 2419-2423.

! M. Haworth and J. McElwain (2008): Hot, dry, wet, cold or toxic? Revisiting the ecological significance of leaf and cuticular micromorphology. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 262: 79–90. See also here.

X. He et al. (2017): Peltaspermalean seed ferns with preserved cuticle from the Upper Triassic Karamay Formation in the Junggar Basin, northwestern China. Abstract, Review of Palaeobotany and Palynology, 247: 68-82. See also here (in PDF).

! J.A. Heredia-Guerrero et al. (2014): Infrared and Raman spectroscopic features of plant cuticles: a review. Free access, Front. Plant Sci., 25.
Note table 2: Definition of semi-quantitative ratios from FTIR and their interpretation for the characterization of fossilized plant cuticles (adapted from Zodrow etal. 2012).

F. Herrera et al. (2018): Exceptionally well-preserved Early Cretaceous leaves of Nilssoniopteris from central Mongolia. Open access, Acta Palaeobotanica, 58: 135–157. See also here.

! A.M. Hetherington and I. Woodward (2003): The role of stomata in sensing and driving environmental change. In PDF, Nature, 424: 901-908.
See also here.

! M.M. Howell et al. (2022): A modified, step-by-step procedure for the gentle bleaching of delicate fossil leaf cuticles. Open access, Fossil Imprint, 78: 445–450.
See also here. "... Previously, the fossil conifer needles from Miocene lignites were consistently destroyed by the use of Schulze’s reagent and produced unusable results with only 5–10% sodium hypochlorite solution. By using the modified weak bleach method given here, large areas of cuticles could be prepared, remained intact, and yielded good diagnostic information on the leaves. ..."

K.R. Hultine and J.D. Marshall (2001): A comparison of three methods for determining the stomatal density of pine needles. In PDF, Journal of Experimental Botany, 52: 369–373. See also here.

A.H. Jahren and N.C. Arens (2009): Prediction of atmospheric δ13CO2 using plant cuticle isolated from fluvial sediment: tests across a gradient in salt content. PDF file, Palaios, 24, 394-401.
Now provided by the Internet Archive´s Wayback Machine.

! M. Javelle et al. (2011): Epidermis: the formation and functions of a fundamental plant tissue. In PDF, New Phytologist, 189: 17-39.

M. Jezek and M.R. Blatt (2017): The membrane transport system of the guard cell and its integration for stomatal dynamics. Free access, Plant Physiology, 174: 487–519.

! T.P. Jones and Nick P. Rowe (eds.), Google Books (some pages are ommitted): Fossil plants and spores: modern techniques. Published by Geological Society, 1999, 396 pages. Excellent! Click: "Preview the book". Go to page 52:
Light microscopy of cuticles (chapter written by H. Kerp and M. Krings).

! G.J. Jordan et al. (2015): Environmental adaptation in stomatal size independent of the effects of genome size. In PDF, New Phytologist, 205: 608-617.

! E.V. Karasev /2013): Formal system of dispersed leaf cuticles of pteridosperms (Peltaspermaceae) from the Permian and Triassic of the Russian platform Paleontological Journal, 47: 335-349. See also here (abstract).

! 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.

Hans Kerp, Palaeobotanical Research Group, University of Münster:
! Plant cuticles and some of their applications in palaeobotany.
This expired link is available through the Internet Archive´s Wayback Machine.

Hans Kerp: The study of fossil gymnosperms by means of cuticular analysis. PALAIOS; 1990; v. 5; no. 6; p. 548-569. See also here (abstract).

G. Kerstiens (1996): Plant cuticles - an integrated functional approach. In PDF, Journal of Experimental Botany.

Gerhard Kerstiens, Institute of Environmental and Natural Sciences, Department of Biological Sciences, Lancaster University: Links to plant surface-related sites.

John W. Kimball, Kimball´s Biology Pages: Gas Exchange in Plants.

J. Konecny, S. Konecny and J. Null, Fossil News, Journal of Avocational Paleontology: The Mazon Creek Nodules.
Still available from the Internet Archive´s Wayback Machine.

! Lenny L.R. Kouwenberg et al. (2007): A new transfer technique to extract and process thin and fragmented fossil cuticle using polyester overlays. Abstract, Review of Palaeobotany and Palynology, 145: 243-248.
See also here (PDF file).

M. Krings et al.(2002): Touch-sensitive glandular trichomes: a mode of defence against herbivorous arthropods in the Carboniferous. PDF file, Evolutionary Ecology Research, 4: 779-786. See also here.

! M. Krings and H. Kerp (1997): An improved method for obtaining large pteridosperm cuticles. In PDF, Review of Palaeobotany and Palynology.

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

E. Kustatscher et al. (2011): Scytophyllum waehneri (Stur) nov. comb., the correct name for Scytophyllum persicum (Schenk) Kilpper, 1975. In PDF, Zitteliana, A 51.

U. Kutschera (2008): The growing outer epidermal wall: Design and physiological role of a composite structure. PDF file, Ann. Bot. 101: 615-621.

U. Kutschera and K.J. Niklas (2007): The epidermal-growth-control theory of stem elongation: An old and a new perspective. PDF file, J. Plant Physiol. 164: 1395-1409.

M.A. Lafuente Diaz et al. (2021): Fourier Transform Infrared Spectroscopy Studies of Cretaceous Gymnosperms from the Santa Cruz Province, Patagonia, Argentina. Abstract, International Journal of Plant Sciences, 182.
Note likewise here (in PDF).
"... The fossils consist of foliar compressions with very well-preserved cuticles, which are chemically characterized by Fourier transform infrared spectroscopy
[...] The compressions [...] probably underwent, during and after diagenesis, a natural oxidation process most likely caused by the recurrent volcanic activity that occurred during the Aptian sedimentation ..."

! T. Linnell (1933); article started on PDF page 21: Zur Morphologie und Systematik triassischer Cycadophyten. II. Über Scytophyllum Bornemann, eine wenig bekannte Cycadophytengattung aus dem Keuper. In PDF, Svensk Botanisk Tidskrift 27: 310–331.
See also here and there.

B.A. Lloyd et al. (2023): CuticleTrace: A toolkit for capturing cell outlines of leaf cuticle with implications for paleoecology and paleoclimatology. Free access, bioRxiv.

B.H. Lomax and W.T. Fraser (2015): Palaeoproxies: botanical monitors and recorders of atmospheric change. In PDF, Palaeontology. See also here (abstract).

B.H. Lomax et al. (2014): Reconstructing relative genome size of vascular plants through geological time. Free access, New Phytologist, 201: 636–644.

! L. Lopez Cavalcante et al. (2023): Analysis of fossil plant cuticles using vibrational spectroscopy: A new preparation protocol. In PDF, Review of Palaeobotany and Palynology, 316.
See also here.
"... alarming changes were caused by the use of Schulze’s solution, which resulted in the addition of both NO2 and (O)NO2 compounds in the cuticle. Consequently, a new protocol using H2CO3, HF, and H2O2 for preparing fossil plant cuticles aimed for chemical analyses is proposed, which provides an effective substitute to the conventional methods ..."

The LoveToKnow Free Online Encyclopedia is based on the eleventh edition of the Encyclopaedia Britannica. Go to:
Websites outdated. Links lead to versions archived by the Internet Archive´s Wayback Machine.

LoveToKnow: The LoveToKnow Free Online Encyclopedia is based on the eleventh edition of the Encyclopaedia Britannica. Go to: Palaeobotany. See also: Preservation.

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

Joyce Macpherson, Memorial University, St. John's, Newfoundland (Canadian Association of Palynologists): Picea Stomata in Lake Sediments. A bibliography. Snapshot taken by the Internet Archive´s Wayback Machine.

M.J.M Martens, R. Aalbers, W.L.P. Janssen, J. van Beurden and E.S. Pierson, General Biology, Katholieke University of Nijmegen, The Netherlands: Virtual Classroom Biology. This extensive site offers educative material about biology, including virtual lessons and lots of illustrations, particularly on cells and tissues. Go to: The microworld of leaves.

L.C.A. Martínez et al. (2020): Studies of the leaf cuticle fine structure of Zuberia papillata (Townrow) Artabe 1990 from Hoyada de Ischigualasto (Upper Triassic), San Juan Province, Argentina. Free access, Review of Palaeobotany and Palynology, 281. See also here (in PDF).

! A.K. Martins et al. (2022): Exceptional preservation of Triassic-Jurassic fossil plants: integrating biosignatures and fossil diagenesis to understand microbial-related iron dynamics. Free access, Lethaia, 55: 1-16. See also here.
Note figure 8: Inferred biogeochemical cycle for the chemical stabilization of iron oxides into goethite in the studied material.
Figure 9: Inferred fossil diagenetic history for the studied fossil plants.
"... there are branches and leaves coated by iron crusts, attributed to the precipitation of iron oxide-oxyhydroxides. Underneath the crusts, the leaves retained minute anatomical features of their epidermal cells and stomatal complexes ..."

N.P. Maslova and A.B. Herman (2015): Approach to Identification of Fossil Angiosperm Leaves: Applicability and Significance of Krassilov´s Morphological System. In PDF, Botanica Pacifica, 4: 103–108.

D. Mauquoy et al. (2010): A protocol for plant macrofossil analysis of peat deposits. PDF file, Mires and Peat, 7.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

! J.C. McElwain et al. (2024): Functional traits of fossil plants. Open access, New Phytologist.
Note figure 2: Examples of fossil plant functional traits.
Figure 4: A ranked list of paleo-functional traits that can be applied to fossil plants.
"What plant remnants have withstood taphonomic filtering, fragmentation, and alteration in their journey to become part of the fossil record provide unique information on how plants functioned in paleo-ecosystems through their traits. Plant traits are measurable morphological, anatomical, physiological, biochemical, or phenological characteristics
[...] We demonstrate how valuable inferences on paleo-ecosystem processes (pollination biology, herbivory), past nutrient cycles, paleobiogeography, paleo-demography (life history), and Earth system history can be derived through the application of paleo-functional traits to fossil plants ..."

! J.C. McElwain and M. Steinthorsdottir (2017): Paleoecology, ploidy, paleoatmospheric composition, and developmental biology: a review of the multiple uses of fossil stomata. Free access, Plant Physiology, 174: 650–664.
See also here.

! Jennifer C. McElwain and William G. Chaloner, Department of Biology, Royal Holloway, University of London, Egham: The fossil cuticle as a skeletal record of environmental change. PDF file, see also here (Abstract), and there.

! S. McLoughlin et al. (2017): The diversity of Australian Mesozoic bennettitopsid reproductive organs. Palaeobio. Palaeoenv., DOI 10.1007/s12549. See also here (in PDF).

! P. Moisan (2012): The study of cuticular and epidermal features in fossil plant impressions using silicone replicas for scanning electron microscopy. In PDF, Palaeontologia Electronica.

Palaeobotany Research Group Münster, Germany:
! Plant cuticles and some of their applications in palaeobotany. An introduction including breathtaking cuticle photomicrographs.
Now provided by the Internet Archive´s Wayback Machine.

Palaeobotany Research Group Münster, Germany:
! History of Palaeozoic Forests, MODES OF PRESERVATION. Link list page with picture rankings. The links give the most direct connections to pictures available on the web.
This expired link is available through the Internet Archive´s Wayback Machine.

! L. Muriale et al. (1996): Fatality due to acute fluoride poisoning following dermal contact with hydrofluoric acid in a palynology laboratory. Free access, Journal of the British Occupational Hygiene Society, 40: 705-710.
! "... The fatality described below highlights the potential for relatively small quantities of concentrated hydrofluoric acid to produce acute systemic toxicity and it is clear that laboratory personnel underestimated the risks ..."

Pyrolysis and macromolecular geochemistry group, Fossil Fuels and Environmental Geochemistry, Newcastle Research Group (NRG), University of Newcastle, Newcastle upon Tyne: The molecular characterization of the very first land plants to appear on the surface of this planet (via wayback link). Abstract.

K.J. Niklas et al. (2017): The evolution of hydrophobic cell wall biopolymers: from algae to angiosperms. Abstract, J. Exp. Bot.

Karl J. Niklas (1981): The Chemistry of Fossil Plants. Abstract, BioScience, 31: 820-825.

! M. Nip et al. (1986): Analysis of modern and fossil plant cuticles by Curie point Py-GC and Curie point Py-GC-MS: recognition of a new, highly aliphatic and resistant biopolymer. In PDF.

N.V. Nosova et al. (2021): Pseudotorellia Florin from the Upper Jurassic–Lower Cretaceous of the Bureya Basin, Russian Far East. Free access, Stratigraphy and Geological Correlation, 29: 434–449.

! N. Nosova et al. (2017): New data on the epidermal structure of the leaves of Podozamites Braun. Abstract, Review of Palaeobotany and Palynology, 238: 88–104. See also here (in PDF).

T.A. Ohsawa et al. (2016): Araucarian leaves and cone scales from the Loreto Formation of Río de Las Minas, Magellan Region, Chile. In PDF, Botany, 94: 805–815. See also here.

M. Özcan et al. (2012): Possible hazardous effects of hydrofluoric acid and recommendations for treatment approach: a review. In PDF, Clinical Oral Investigations, 16: 15–23. See also here.

J. Pittermann (2010): The evolution of water transport in plants: an integrated approach. In PDF, Geobiology.
See also here.

M. Pole (2008): The record of Araucariaceae macrofossils in New Zealand. Free access, Alcheringa, 32: 405–426.

Mike Pole, Queensland Herbarium, Toowong, Australia: Early Eocene Dispersed Cuticles and Mangrove to Rainforest Vegetation at Strahan-Regatta Point, Tasmania. Paleontologia Electronica 2007, 10 (3).

! C. Pott et al. (2012): Trichomes on the leaves of Anomozamites villosus sp. nov. (Bennettitales) from the Daohugou beds (Middle Jurassic), Inner Mongolia, China: Mechanical defence against herbivorous arthropods. In PDF, Review of Palaeobotany and Palynology, 169: 48-60.
See also here.

C. Pott et al. (2008): The Carnian (Late Triassic) flora from Lunz in Lower Austria: Paleoecological considerations. In PDF, Palaeoworld, 17: 172-182. See also here.

! C. Pott and H. Kerp (2008): Mikroskopische Untersuchungsmethoden an fossilen Pflanzenabdrücken. In PDF. Der Präparator.

! C. Pott (2007): Cuticular analysis of gymnosperm foliage from the Carnian (Upper Triassic) of Lunz, Lower Austria. In PDF, Thesis, Westfälische Wilhelms-Universität Münster, Germany.

J.A. Raven (2002): Selection pressures on stomatal evolution. PDF file, New Phytologist.

H. Renault et al. (2017): A phenol-enriched cuticle is ancestral to lignin evolution in land plants. Nat. Commun. 8.

K.S. Renzaglia et al. (2017): Hornwort stomata: architecture and fate shared with 400 million year old fossil plants without leaves. Free access, Plant Physiology, 177: 788–797.

! G.J. Retallack (2001): A 300-million-year record of atmospheric carbon dioxide from fossil plant cuticles. In PDF, Nature.
This expired link is available through the Internet Archive´s Wayback Machine. See also:
Supplementary Information for "A 300-million-year record of atmospheric carbon dioxide from fossil plant cuticles" Nature, V411, 287. They are measurements of stomatal index from fossil and living plants. Part 1 has reliable data, and Part 2 has data deemed statistically inadequate from a rarefaction analysis. Abbreviations include SI (stomatal index), Nf (number of fragments counted), Ns (number of stomates counted), Ne (number of epidermal cells counted), and Ma (millions of years ago).

M. Riederer and L. Schreiber et al. (2001): Protecting against water loss: analysis of the barrier properties of plant cuticles. Journal of experimental botany, 52: 2023-2032. In PDF, see also here.

Markus Riederer, Julius von Sachs Institut, Würzburg: Stoffaustausch über pflanzliche Grenzflächen (in German). Research about plant cuticles.
Now recovered from the Internet Archive´s Wayback Machine.

M. Riederer and L. Schreiber et al. (1995): Waxes: the transport barriers of plant cuticles. PDF file, in: R.J. Hamilton (ed.). Waxes: Chemistry, Molecular Biology and Functions. The Oily Press, West Ferry, Dundee, Scotland.

S.J. Rogerson et al. (1976): An improved preparation technique for identification of plant cuticle in animal faeces. In PDF, New Zealand Journal of Botany, 14: 117-119.

Anita Roth-Nebelsick (2007): Computer-based Studies of Diffusion through Stomata of Different Architecture. PDF file, Ann. Bot., 100: 23-32. See also here.

P.J. Rudall and R.M. Bateman (2019): Leaf surface development and the plant fossil record: stomatal patterning in Bennettitales. Abstract, Biological Reviews.
"... Fossil bennettites – even purely vegetative material – can be readily identified by a combination of epidermal features, including distinctive cuticular guard-cell thickenings, lobed abaxial epidermal cells (‘puzzle cells’), transverse orientation of stomata perpendicular to the leaf axis, and a pair of lateral subsidiary cells adjacent to each guard-cell pair (termed paracytic stomata). ..."

P.J. Rudall et al. (2017): Evolution and development of monocot stomata. In PDF, American journal of botany, 104: 1122-1141.

E.-M. Sadowski et al. (2017): Conifers of the "Baltic amber forest" and their palaeoecological significance. In PDF, Stapfia, 106. See also here.
Note Fig. 1: Terminology of the stomata morphology.

P. Sarkar et al. (2009): Plant cell walls throughout evolution: towards a molecular understanding of their design principles. In PDF, Journal of Experimental Botany, 60: 3615–3635. See also here.

E. Schneebeli-Hermann et al. (2014): Vegetation history across the Permian–Triassic boundary in Pakistan (Amb section, Salt Range). Gondwana research, 27: 911-924.
See also here, and there (in PDF).

A.C. Scott and G. Rex (1985): The formation and significance of Carboniferous coal balls. PDF file, Philosophical Transactions of the Royal Society London, B, 311: 123-137.
See also here, and there.

M. Seale (2020): The Fat of the Land: Cuticle Formation in Terrestrial Plants. Free access, Plant Physiology, 184: 1622–1624.

Z. Simunek and J. Haldovský (2015): Contribution to the knowledge of Cordaites species from the Kladno-Rakovník Basin, Middle Pennsylvanian (Bolsovian), Czech Republic. In PDF, Geologia Croatica.

Z. Simunek et al. (2009): Cordaites borassifolius (Sternberg) Unger (Cordaitales) from the Radnice Basin (Bolsovian, Czech Republic). PDF file, Bulletin of Geosciences, 84: 301-336.

! M. Slodownik et al. (2023): Komlopteris: A persistent lineage of post-Triassic corystosperms in Gondwana. Free access, Review of Palaeobotany and Palynology, 317.
Note figure 1A: Geochronological scale indicating the range of Southern Hemisphere Komlopteris species.
"... Komlopteris is a genus that includes the youngest representative of the so-called ‘seed ferns’
[...] we review the representatives of Komlopteris from Gondwana, emend the genus, establish three new species, and propose five new combinations based on macro-morphological traits ..."

W.K. Soh et al. (2017): Palaeo leaf economics reveal a shift in ecosystem function associated with the end-Triassic mass extinction event. Abstract, Nature plants, 3. See also here (supplementary information) and there (corrigendum, in PDF).

! R.A. Spicer (1989): Physiological characteristics of land plants in relation to environment through time. In PDF, Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 80.
See also here.

! R.A. Spicer (1977): The pre-depositional formation of some leaf impressions. PDF file, Palaeontology, 20: 907–912.
This expired link is now available through the Internet Archive´s Wayback Machine.

B. Artur Stankiewicz et al. (1998): Chemical preservation of plants and insects in natural resins. PDF file, Proc. R. Soc. Lond. B, 265: 641-647. See also here.

B.A. Stankiewicz et al. (1998): Molecular taphonomy of arthropod and plant cuticles from the Carboniferous of North America: implications for the origin of kerogen. In PDF, Journal of the Geological Society, 155: 453-462.
See also here.

M. Steinthorsdottir et al. (2018): Cuticle surfaces of fossil plants as a potential proxy for volcanic SO2 emissions: observations from the Triassic–Jurassic transition of East Greenland. In PDF, Palaeobiodiversity and Palaeoenvironments, 98: 49–69. See also here.

! M. Steinthorsdottir et al. (2011): Extremely elevated CO2 concentrations at the Triassic/Jurassic boundary. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 308: 418-432.
See also here.
"... The final results indicate that pre-TJB (Rhaetian), the CO2 concentration was approximately 1000 ppm, that it started to rise steeply pre-boundary and had doubled to around 2000–2500 ppm at the TJB. The CO2 concentration then remained elevated for some time post-boundary, before returning to pre-TJB levels in the Hettangian. ..."

E.J. Stevens et al. (1987): Procedure for Fecal Cuticle Analysis of Herbivore Diets. PDF file.

! 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.

Taylor S. Feild, Maciej A. Zwieniecki, Michael J. Donoghue, and N. Michele Holbrook: Stomatal plugs of Drimys winteri (Winteraceae) protect leaves from mist but not drought. PNAS, 1998 95: 14256-14259.

! G.R. Upchurch Jr. (1995): Dispersed angiosperm cuticles: Their history, preparation, and application to the rise of angiosperms in Cretaceous and Paleocene coals, southern western interior of North America. In PDF, International Journal of Coal Geology, 28: 161-227. See also here.

! G.A. Upchurch Jr. (1989): Dispersed angiosperm cuticles. In PDF, Notes for a Workshop on the Study of Fragmentary Plant Remains.

G.R. Upchurch Jr. (1984): Cuticle evolution in Early Cretaceous angiosperms from the Potomac Group of Virginia and Maryland. PDF file, Annals of the Missouri Botanical Garden. A version archived by Internet Archive Wayback Machine.

G.R. Upchurch Jr. (1984): The cuticular anatomy of early angiosperm leaves from the Lower Cretaceous Potomac Group of Virginia and Maryland, Part 1, Zone 1 leaves. Snapshot taken by the Internet Archive´s Wayback Machine. PDF file, American Journal of Botany 71: 192-202.

M.A. Urban et al. (2018): Cuticle and subsurface ornamentation of intact plant leaf epidermis under confocal and superresolution microscopy. In PDF, Microsc. Res. Tech. 81, 129–140.
See also here and there.

! V. Vajda et al. (2017): Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships. In PDF, Nature Ecology & Evolution. See also here and there.

! P.F. van Bergen et al. (1995): Resistant biomacromolecules in the fossil record. Abstract, Acta botanica neerlandica. See also here (in PDF).

! A. Vatén and D.C. Bergmann (2012): Mechanisms of stomatal development: an evolutionary view. In PDF, EvoDevo, 3.

! H. Visscher (1993): Links with the past in the plant world: cuticles as recorders of diversity, kerogen formation and palaeoatmospheric CO2-level. In PDF, The Palaeobotanist.

L. Wang and Q. Leng (2011): A new method to prepare clean cuticular membrane from fossil leaves with thin and fragile cuticles. In PDF, Science China Earth Sciences, 54: 223-227. See also here.

Y. Wang et al. (2005): Cuticular anatomy of Sphenobaiera huangii (Ginkgoales) from the lower Jurassic of Hubei, China. In PDF, American Journal of Botany, 92: 709-721.

Jing-Ke Weng and Clint Chapple (2010): The origin and evolution of lignin biosynthesis. New Phytologist, 187: 273-285.

Wikipedia, the free encyclopedia Plant cuticle.

! J.P. Wilson et al. (2020): Carboniferous plant physiology breaks the mold. Free access, New Phytologist.

! J.P. Wilson et al. (2017): Dynamic carboniferous tropical forests: new views of plant function and potential for physiological forcing of climate. Free access, New Phytologist, 215: 1333–1353.

M.J. Wooller (2002): Fossil grass cuticles from lacustrine sediments: a review of methods applicable to the analysis of tropical African lake cores. PDF file, The Holocene.

X.-J. Yang et al. (2009): Leaf cuticle ultrastructure of Pseudofrenelopsis dalatzensis (Chow et Tsao) Cao ex Zhou (Cheirolepidiaceae) from the Lower Cretaceous Dalazi Formation of Jilin, China. PDF file, Review of Palaeobotany and Palynology, 153: 8-18.
The link is to a version archived by the Internet Archive´s Wayback Machine.
See also here.

! T.H. Yeats and J.K.C. Rose (2013): The formation and function of plant cuticles. In PDF, Plant physiology, 163: 5–20. See also here.

B. Zhang et al. (2024): Numerical taxonomy and genus-species identification of Czekanowskiales in China based on machine learning. Free access, Palaeontologia Electronica, 27.
"... accurate identification of Czekanowskiales fossils is difficult due to the similarities in some macroscopic and cuticular patterns among different genera and species
[...] This study focused on the numerical taxonomy and identification of Czekanowskiales at the generic and species levels using cluster analysis, trait selection, and supervised learning methods for machine learning ..."

M. Zhao et al. (2015): Anomozamites (Bennettitales) from Middle Jurassic Haifanggou Formation, western Liaoning, China. In PDF, Global Geology, 18: 75-87.

Carl Zimmer (Carl Zimmer writes the monthly essay in the US magazine Natural History, having inherited this position from Stephen Jay Gould): High and dry. Stomatal apparatus permitting plants to become trees. A version archived by Internet Archive Wayback Machine.

E.L. Zodrow and J.A. D'angelo (2013): Digital compression maps: an improved method for studying Carboniferous foliage. In PDF, Atlantic Geology, 49. See also here and there.
"... The image of any freed frond segment of compression foliage that has been reprocessed digitally to represent its original structure is called a compression map. ..."

E.L. Zodrow et al. (2010): Phytochemistry of the fossilized-cuticle frond Macroneuropteris macrophylla (Pennsylvanian seed fern, Canada). Abstract, International Journal of Coal Geology, 84: 71-82.

E. Zodrow and M. Mastalerz (2009): A proposed origin for fossilized Pennsylvanian plant cuticles by pyrite oxidation (Sydney Coalfield, Nova Scotia, Canada). PDF file, Bulletin of Geosciences, 84: 227-240.

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