Home / Palaeoclimate / Stomatal Density

Karsten Alfermann (2001): Untersuchungen zur Anpassung des Photosyntheseapparates Höherer Pflanzen bei Wachstum unter einem erhöhtem CO₂-Partialdruck von 700 ppm. PDF file (9.3 MB), in German. Dissertation, Universität Bielefeld.

R. Barclay, J. McElwain, D. Dilcher and B. Sageman (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.

A. Baresch et al. (2019): Competition for epidermal space in the evolution of leaves with high physiological rates. In PDF, New Phytologist, 221: 628-639. See also here.

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.

D.J. Beerling et al. (1998): Stomatal responses of the "living fossil" Ginkgo biloba L. to changes in atmospheric CO₂ concentrations. PDF file, Journal of Experimental Botany, 49: 1603-1607.

David Beerling, White Rose Palaeobiology Group, UK: Atmospheric CO₂ and climate change during the Permo-Carboniferous glaciation inferred from fossil plants. Project description. See also here (Low atmospheric CO₂ levels during the Permo- Carboniferous glaciation inferred from fossil lycopsidsPDF file, in PDF).
These expired links are available through the Internet Archive´s Wayback Machine.

Beerling DJ, Lake JA, Berner RA, Hickey LJ, Taylor DW, Royer DL. 2002: Carbon isotope evidence implying high O₂/CO₂ ratios in the Permo-Carboniferous atmosphere. PDF file, from Geochimica et Cosmochimica Acta, 66: 3757-3767.
Snapshot taken by the Internet Archive´s Wayback Machine.

D. J. Beerling, B. H. Lomax, D. L. Royer, G. R. Upchurch, Jr., and L. R. Kump: An atmospheric pCO₂ reconstruction across the Cretaceous-Tertiary boundary from leaf megafossils. The National Academy of Sciences, PNAS 2002 99: 7836-7840.

D. J. Beerling Department of Animal and Plant Sciences, University of Sheffield, and D. L. Royer, Department of Geology and Geophysics, Yale University, New Haven: Reading a CO₂ signal from fossil stomata. Research review, PDF file (New Phytologist, 2002; 153: 387– 397).
This expired link is available through the Internet Archive´s Wayback Machine.

Robert A. Berner, Geology and Geophysics, Yale University, New Haven, Connecticut: The Rise of Plants and Their Effect on Weathering and Atmospheric CO₂ (now via wayback archive). See also here, and there.

Robert A. Berner, Department of Geology and Geophysics, Yale University, New Haven, CT: Atmospheric oxygen over Phanerozoic time. PNAS, Vol. 96, Issue 20, 10955-10957, September 28, 1999.

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).

! L.T.&xnbsp;Bertolino et al. (2019): Impact of Stomatal Density and Morphology on Water-Use Efficiency in a Changing World. Free access, Front. Plant Sci. 10:225. doi: 10.3389/fpls.2019.00225.

M.R. Blatt et al. (2024): Photosynthesis and the stomatal nexus, past, present and future. Open access, Plant, Cell & Environment.

! Bert Bolin, Egon T. Degens, Stephan Kempe, and Pieter Ketner 1979 (illustrated HTML at icsu-scope.org, SCOPE, The Scientific Committee On Problems of the Environment): The Global Carbon Cycle. This expired link is available through the Internet Archive´s Wayback Machine.
For instance: The Possible Effects of Increased CO₂ on Photosynthesis. by J. Goudriaan and Jr. G. L. Ajtay.

! N.R. Bonis (2010), Laboratory of Palaeobotany and Palynology, Palaeoecology Institute of Environmental Biology, Department of Biology, Utrecht University: Palaeoenvironmental changes and vegetation history during the Triassic-Jurassic transition. PDF file (7.7 MB), LPP Contribution Series No. 29. Seven research reports (chapters) in this thesis, see especially chapter 7 (with W.M. Kürschner):
! Vegetation history, diversity patterns, and climate change across the Triassic-Jurassic boundary (PDF page 140).
Provided by the Internet Archive´s Wayback Machine.
See also here.

C.K. Boyce and M.A. Zwieniecki (2018): The prospects for constraining productivity through time with the whole-plant physiology of fossils. Open access, New Phytologist.

C.K. Boyce and A.B. Leslie (2012): The Paleontological Context of Angiosperm Vegetative Evolution. In PDF, International Journal of Plant Sciences, 173: 561–568.
See also here.
"... a survey of the fossil record demonstrates that most anatomical traits that are now unique to the angiosperms were more broadly distributed among extinct lineages.
[...] Of all the various vegetative morphological traits that have been traditionally linked to angiosperm success, only the evolution of very high leaf vein densities appears to be truly unique to the angiosperms. ..."

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).

T.J. Brodribb et al. (2007): Leaf Maximum Photosynthetic Rate and Venation Are Linked by Hydraulics. Open access, Plant Physiology, 144: 1890-1898.

! 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.

Steve Case, University of Kansas Lawrence (page hosted by Access Excellence): Leaf Stomata as Bioindicators of Environmental Change.

! Center for the Study of Carbon Dioxide and Global Change: Are Physical Properties of Stomata Unresponsive to CO₂? Provided by the Internet Archive´s Wayback Machine.
Stomatal density (SD; the number of stomata per unit leaf area) and stomatal index (SI; the number of stomata divided by the sum of the numbers of stomatal and epidermal cells) are often used as proxies of past atmospheric CO₂ concentrations. The evidence for the validity of this technique is said by Reid et al. to rest primarily upon paleontological data and growth chamber studies. Hence, they sought to broaden the experimental basis for the protocol by adding field studies to the mix of evidence supporting it.

! Center for the Study of Carbon Dioxide and Global Change, Tempe, AZ. This center was created to disseminate factual reports and sound commentary on new developments in the world-wide scientific quest to determine the climatic and biological consequences of the ongoing rise in the air's CO₂ content. Go to: Stomatal Density. Directory of articles, e.g. Stomatal Frequency Responses of Conifer Needles to Atmospheric CO₂ Enrichment.
These expired links are available through the Internet Archive´s Wayback Machine.

G. Chen et al. (2024): Stomatal evolution and plant adaptation to future climate. Open access, Plant Cell Environ.
"Global climate change is affecting plant photosynthesis and transpiration processes, as well as increasing weather extremes
[...] novel stomatal development specific genes were acquired during plant evolution, whereas genes regulating stomatal movement, especially cell signaling pathways, were inherited ancestrally and co-opted by dynamic functional differentiation ..."

! CLAMP Online (Climate Leaf Analysis Multivarite Program). This site is the result of an ongoing collaboration between the Institute of Botany, Chinese Academy of Sciences, Beijing, and the Open University UK.
How you can use foliar physiognomy (leaf architecture) to determine ancient climates from fossil leaves or explore the relationship that exists between leaf form and climate. CLAMP is a multivariate statistical technique that decodes the climatic signal inherent in the physiognomy of leaves of woody dicotyledonous plants. See especially:
! Teaching Materials.
Older CLAMP websites are available through the Internet Archive´s Wayback Machine:
Robert A. Spicer, The Warm Earth Environmental Systems Research Group: Plant Fossils as Climatic Indicators. Go to: Climate Leaf Analysis Multivariate Programe (CLAMP). An introduction to the use of leaf architecture for determining past climatic conditions.

! 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.

Tom Clarke, Nature scienceupdate, 17 May 2001: Climatologists pore over past. Fossil leaves tell us about the air the dinosaurs breathed.
Still provided by the Internet Archive´s Wayback Machine.

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. 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.
Website outdated, a version archived by the Internet Archive 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 ..."

UCD Plant Palaeoecology and Palaeobiology Group, Dublin, Ireland:
OXYEVOL: The role of atmospheric oxygen in plant evolution over the past 400 million years.
The aim of the project is to identify how changes in atmospheric O₂ and CO₂ concentration influence the timing of key evolutionary innovations and shifts in ecological dominance/success of various plant groups throughout geological time.

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.

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.

! 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 ..."

C. Elliott-Kingston et al. (2016): Does size matter? Atmospheric CO₂ may be a stronger driver of stomatal closing rate than stomatal size in taxa that diversified under low CO₂. In PDF, Front Plant Sci., 7. See also here and there.

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.

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).

A. Fangmeier & H.-J. Jäger, Institut für Pflanzenökologie der Justus-Liebig-Universität Gießen: 4.2 Wirkungen erhöhter CO₂-Konzentrationen. PDF file (in German), from Guderian, R. (ed.): Handbuch der Umweltveränderungen und Ökotoxikologie; Band 2a: Terrestrische Ökosysteme. Berlin: Springer, 2001.

Ben Fletcher, Department of Animal and Plant Sciences, University of Sheffield: How the atmosphere affects plants. See also: The role of stomata in the early evolution of land plants.
Snapshots provided by the Internet Archive´s Wayback Machine.

! J.P. Fortin and W.E. Friedman (2024): A stomate by any other name? The open question of hornwort gametophytic pores, their homology, and implications for the evolution of stomates. Free access, New Phytologist. https://doi.org/10.1111/nph.20094.
"... We explore the occurrence and diverse functions of stomates throughout the evolutionary history and diversity of extinct and extant embryophytes. We then address arguments for and against homology between known sporophyte- and gametophyte-borne stomates and HGPs [hornwort gametophytic pores] and conclude that there is little to no evidence that contradicts the hypothesis of homology ..."

P.J. Franks and D.L. Royer (2017): Comment on "Was atmospheric CO₂ capped at 1000ppm over the past 300millionyears?" by McElwain JC et al. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 472: 256–259. See also here.

P.J. Franks et al. (2012): Megacycles of atmospheric carbon dioxide concentration correlate with fossil plant genome size. In PDF, Phil. Trans. R. Soc. B, 367: 556-564.
See also here.

! Peter J. Franks and David J. Beerling (2009): Maximum leaf conductance driven by CO₂ effects on stomatal size and density over geologic time. PDF file, PNAS, 106: 10343-10347.

Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine: PLANTS AS KEYS TO PAST CLIMATIC CONDITIONS.
Still available through the Internet Archive´s Wayback Machine.
They are spatially-fixed adapted to atmospheric and substrate conditions. They are tightly constrained by the climatic regime under which they grow. See also Is it possible to retrodict? Fossils as environmental indicators.

! A. Gessler et al. (2014): Stable isotopes in tree rings: towards a mechanistic understanding of isotope fractionation and mixing processes from the leaves to the wood. Free access, Tree Physiology, 34: 796–818.
Note figure 1: Overview of the different processes influencing the carbon (a) and oxygen (b) isotope signature, from primary sources (CO₂ and H₂O, respectively) to tree-ring cellulose, going through different organic and inorganic pools.

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.

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

M. Haworth et al. (2011): Cycads show no stomatal-density and index response to elevated carbon dioxide and subambient oxygen. Abstract, Australian Journal of Botany.

Matthew Haworth et al. (2010): Differences in the response sensitivity of stomatal index to atmospheric CO2 among four genera of Cupressaceae conifers. PDF file, Ann. Bot., 105: 411-418.

! 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.

! 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.

K.E. Hill et al. (2019): Pinnule and Stomatal Size and Stomatal Density of Living and Fossil Bowenia and Eobowenia Specimens Give Insight into Physiology during Cretaceous and Eocene Paleoclimates. Free access, Int. J. Plant Sci., 180: 323–336.

The Hindu, India: Fossil leaves reveal Earth's history.
Still available via Internet Archive Wayback Machine.

J. Hošek et al. (2024): Hot spring oases in the periglacial desert as the Last Glacial Maximum refugia for temperate trees in Central Europe. Free access, Science Advances, 10, eado6611.
Note figure 1: European paleoenvironments during the LPG (~28 to 14.7 ka).

R.A. Houghton, Woods Hole Research Center,MA: The Contemporary Carbon Cycle. (PDF file). A sample chapter of Volume 8. Biogeochemistry (William H. Schlesinger), Treatise on Geochemistry.

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.

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.

! 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.

P. Kenrick (2001): Turning over a new leaf. PDF file, Nature, 410: 309-310. This expired link is available through the Internet Archive´s Wayback Machine.

Richard A. Kerr, Science magazine, April 2005: Gasping for Air in the Permian. Thin air may have forced animals down from higher latitudes 250 million years ago, crowding them into the lowlands and possibly helping along the largest extinction in the history of the planet, according to a study of Science.

Hans Kerp, Forschungsstelle für Paläobotanik, Geologisch-Paläontologisches Institut, Westfälische Wilhelms-Universität, Münster, Germany: Palaeobotany (Communications arising): Atmospheric CO₂ from fossil plant cuticles. Abstract, Nature 415, 38 (2002). See also here

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

Lenny L.R. Kouwenberg et al. (2007): Stomatal Frequency Change Over Altitudinal Gradients: Prospects for Paleoaltimetry. PDF file, Reviews in Mineralogy & Geochemistry, 66: 215-241.
See also here.

L. Kouwenberg et al. (2005): Atmospheric CO₂ fluctuations during the last millennium reconstructed by stomatal frequency analysis of Tsuga heterophylla needles. PDF file, Geology, 33: 33-36.
See likewise here.

W.M. Kürschner (2001): Leaf sensor for CO₂ in deep time. In PDF, News and views, Nature, 411: 247-48.
See also here.

W.M. Kürschner et al. (1996): Oak leaves as biosensors of late Neogene and early Pleistocene paleoatmospheric CO₂ concentrations. In PDF, Marine Micropaleontology, 27: 299-312.
See also here.

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.

! A.D.B. Leakey and J.A. Lau (2012): Evolutionary context for understanding and manipulating plant responses to past, present and future atmospheric [CO₂]. Phil. Trans. R. Soc. B, 367: 613-629. See als here (in PDF).

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.

! J. Loranger and B. Shipley (2010): Interspecific covariation between stomatal density and other functional leaf traits in a local flora. Abstract, Botany, 88: 30-38.

B.A. Lloyd et al. (2024): CuticleTrace: A toolkit for capturing cell outlines from leaf cuticle with implications for paleoecology and paleoclimatology. Free access, Applications in Plant Sciences, 12.

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.

! 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.

! J.C. McElwain et al. (2016): Was atmospheric CO₂ capped at 1000 ppm over the past 300 million years? In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 441: 653–658. See also here.

! J.C McElwain et al. (2015): Using modern plant trait relationships between observed and theoretical maximum stomatal conductance and vein density to examine patterns of plant macroevolution. New Phytologist, 209: 94-103.

Jennifer C. McElwain, UCD Earth Systems Institute, Dublin: Climate change and mass extinction: What can we learn from 200 million year old plants? PDF file.
Provided by the Internet Archive´s Wayback Machine.

Jennifer C. McElwain, Department of Geology, The Field Museum, Chicago: A novel climate-independent method for estimating paleo-elevation from fossil plants. Abstract; Botany 2001 "Plants and People" August 12 - 16, 2001, Albuquerque.

! J.C. McElwain and W.G. Chaloner (1995): Stomatal density and index of fossil plants track atmospheric carbon dioxide in the Palaeozoic. PDF file, Annals of Botany, 76: 389-395.

! 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.

Jennifer C. McElwain, Jessica Wade-Murphy and Stephen P. Hesselbo: Changes in carbon dioxide during an oceanic anoxic event linked to intrusion into Gondwana coals. Abstract, Nature 435: 479-482; May 2005. Using the stomatal index method. Although multiple forcing factors may have contributed to the abrupt spike in atmospheric CO₂, the authors suggest that the likely dominant forcing factor was oxidation of methane gas generated by subsurface thermal metamorphism of organic-rich late Permian and late Triassic coal bearing strata during magmatic intrusion of the Karoo-Ferrar large igneous province of southern Gondwana.

J.N. Milligan et al. (2022): Moderate to elevated atmospheric CO₂ during the early Paleocene recorded by Platanites leaves of the San Juan Basin, New Mexico. In PDF, Paleoceanography and Paleoclimatology, 37, e2021PA004408.
See also here.

mysan.de/international (a Montrasio and Illing GdbR News Portal): Holy Grail of Geology Found: Measuring Elevation Over Geological Eras. New method (counting the stomata on leaves of fossil plants) to measure ancient land elevation developed by Field Museum scientist. See also here (Der Spiegel, in German).

New Phytologist, Forum: Remote control – cell and organ communication within plants. PDF file.

Larry O'Hanlon, Discovery News: Ancient Fossil Fuels Caused Jurassic Warming. The carbon dioxide level and the stomata method. Provided by the Internet Archive´s Wayback Machine.

Larry Orr, Photosynthesis Center, Arizona State University, Tempe, AZ: What is Photosynthesis? Links (with annotations) to articles that discuss photosynthesis at varying degrees of complexity.

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.

Osborne, C.P. & Beerling, D.J. (2002): A process-based model of conifer structure and function with special emphasis on leaf lifespan.. Global Biogeochemical Cycles.
The link is to a version archived by the Internet Archive´s Wayback Machine.

J.L. Payne et al. (2006): The Pattern and Timing of Biotic Recovery from the End-Permian Extinction on the Great Bank of Guizhou, Guizhou Province, China. In PDF, Palaios, 21: 63-85.
This expired link is now available through the Internet Archive´s Wayback Machine.

! N. Pérez-Harguindeguy et al. (2013): New handbook for standardised measurement of plant functional traits worldwide. In PDF, Australian Journal of Botany, 61: 167-234.

Sara Pratt, Geotimes: Reaching past heights. About methods calculating paleoelevations.

J.A. Raven (2017): Evolution and palaeophysiology of the vascular system and other means of long-distance transport. In PDF, Phil. Trans. R. Soc. B, 373: 20160497.

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

J. Read and A. Stokes (2006): Plant biomechanics in an ecological context. Free access, American Journal of Botany, 93: 1546-1565.

Chantal D. Reid, Robert B. Jackson (Department of Biology, Duke University, Durham, NC), and Joy K. Ward Department of Biology, University of Utah, Salt Lake City, UT: Carbon dioxide as a selective agent for stomatal density. Abstract.

! G.J. Retallack (2001): A 300-million-year record of atmospheric carbon dioxide from fossil plant cuticles. In PDF, Nature. 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).

! G.J. Retallack (2002):&xnbsp; Carbon dioxide and climate over the past 300 Myr. In PDF, Phil. Trans. R. Soc. Lond., A, 360: 659–673.
See also here.

G.J.Retallack, University of Oregon, Eugene: Soils and Global Change in the Carbon Cycle over Geological Time (PDF file).

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.

Sue Rigby, Geology, Geophysics, Environmental Geoscience, Grant Institute, University of Edinburgh: COURSE MATERIALS. Go to: GEP COURSE MATERIALS, Lecture 8: Impacts of life on the planet. PDF file.

Daniel H. Rothman, Department of Earth, Atmospheric, and Planetary Sciences, Massachusetts Institute of Technology, Cambridge, MA: Global biodiversity and the ancient carbon cycle. Proc. Natl. Acad. Sci. USA, Vol. 98, Issue 8, 4305-4310, April 10, 2001.

Daniel H. Rothman, Department of Earth, Atmospheric, and Planetary Sciences, Massachusetts Institute of Technology, Cambridge, MA: Atmospheric carbon dioxide levels for the last 500 million years. Proc. Natl. Acad. Sci. USA, Vol. 99, Issue 7, 4167-4171, April 2, 2002.

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

! A. Roth-Nebelsick et al. (2001): Evolution and Function of Leaf Venation Architecture: A Review. PDF file, Annals of Botany 87: 553-566. See also here.

D.L. Royer et al. (2002): High CO₂ increases the freezing sensitivity of plants: implications for paleoclimatic reconstructions from fossil floras.. In PDF, Geology, 30: 963-966.
The link is to a version archived by the Internet Archive´s Wayback Machine.

D.L. Royer (2001): Stomatal density and stomatal index as indicators of paleoatmospheric CO2 concentration. PDF file, from Review of Palaeobotany and Palynology 114 (2001) 1-28.
Snapshot provided by the Internet Archive´s Wayback Machine.

! D.L. Royer et al. (2001): Phanerozoic atmospheric CO₂ change: evaluating geochemical and paleobiological approaches. In PDF, Earth-Science Reviews, 54: 349-392.
See also here.

D.L. Royer, Department of Geosciences, Pennsylvania State University, University Park, PA: Using stomatal distributions to reconstruct ancient levels of atmospheric CO₂.
Now recovered from the Internet Archive´s Wayback Machine.

L. Sack and T.N. Buckley (2016): The developmental basis of stomatal density and flux. Free access, Plant physiology, 171: 2358–2363.

L. Santasalo (2013): The Jurassic extinction events and its relation to CO₂ levels in the atmosphere: a case study on Early Jurassic fossil leaves. In PDF, Bachelor´s thesis, Department of Geology, Lund University, Sweden.

Science and Plants for Schools (SAPS).
SAPS creates opportunities for teachers, technicians and students to find out more about plants and to become more interested in plant science.
Measuring Stomatal Density.

ScienceDaily Magazine: Fossil Plants' Ties To Ancient Carbon Redefined.

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

Laura Serna (2008): Drawing the future: Stomatal response to CO₂ levels. PDF file, Plant Signaling and Behavior 3: 214-217. See also here.

M. Slodownik et al. (2021): Fossil seed fern Lepidopteris ottonis from Sweden records increasing CO₂ concentration during the end-Triassic extinction event. Open access, Palaeogeography, Palaeoclimatology, Palaeoecology, 564. See also here (in PDF).

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).

Nancy E. Spaulding & Samuel N. Namowitz (McDougal Littell): Exploring Earth. The investigations and visualizations on this site were designed to accompany Earth Science, a high school textbook. The Web site was developed by TERC, a non-profit educational research and development firm in collaboration with McDougal Littell. Funding was provided by the National Science Foundation. Go to: The greenhouse effect and global warming.

! R.A. Spicer (1992): Fossils as Environmental Indicators, Climate from Plants. PDF file.
Now recovered from the Internet Archive´s Wayback Machine.

R.A. Stein et al (2024): Comparing Methodologies for Stomatal Analyses in the Context of Elevated Modern CO₂. Open access, Life, 14. https://doi.org/10.3390/life14010078

! M. Steinthorsdottir et al. (2022): Key traits of living fossil Ginkgo biloba are highly variable but not influenced by climate – Implications for palaeo-pCO₂ reconstructions and climate sensitivity. In PDF, Global and Planetary Change, 211. See also here.

! M. Steinthorsdottir et al. (2021): Searching for a nearest living equivalent for Bennettitales: a promising extinct plant group for stomatal proxy reconstructions of Mesozoic _pCO₂. Open access, GFF, 143: 190-201.

M. Steinthorsdottir and V. Vajda (2013): Early Jurassic (late Pliensbachian) CO₂ concentrations based on stomatal analysis of fossil conifer leaves from eastern Australia. In PDF, Gondwana Research.

Teaching Issues and Experiments in Ecology (TIEE): TIEE is a project of the Education and Human Resources Committee of the Ecological Society of America. Go to: Environmental Correlates with Leaf Stomata Density. Detailed Description of the Experiment. See also here.

Dieter Uhl and Hans Kerp, Forschungsstelle für Paläobotanik, Westfälische Wilhelms-Universität, Münster: Stomatal-density and -index in Upper Permian conifers. Abstract, The International Plant Taphonomy Meeting 2002, Bonn, Goldfuss Museum, Institute of Paleontology, Germany.

Henk Visscher, Utrecht University: Fossil leaves as biosensors of paleoatmospheric CO₂ levels. Abstract (RTF file) of the 2000 lecture series of Petroleum Geologische Kring.
Snapshot provided by the Internet Archive´s Wayback Machine.

R.V. Vofély et al. (2019): Of puzzles and pavements: a quantitative exploration of leaf epidermal cell shape. Free access, New Phytologist, 221: 540-552.

Friederike Wagner, Raimond Below, Pim De Klerk, David L. Dilcher, Hans Joosten, Wolfram M. Kürschner, and Henk Visscher: A natural experiment on plant acclimation: Lifetime stomatal frequency response of an individual tree to annual atmospheric CO₂ increase. The National Academy of Sciences, PNAS 1996 93: 11705-11708.

The White Rose Palaeobiology Group. This is a collaborative initiative between the Universities of Sheffield and Leeds. Go to: Atmospheric CO₂ - the key to early leaf evolution.
These expired links are now available through the Internet Archive´s Wayback Machine.

! 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.

SanPing XIE et al. (2009): Altitudinal variation in Ginkgo leaf characters: Clues to paleoelevation reconstruction. PDF file, Science in China Series D: Earth Sciences, 52: 2040-2046.
"The results show that leaf area, petiole length, and stomatal parameters have no obvious linear relationship with altitude (...). The results also suggest that the differences in stomatal density and stomatal index between sun and shade leaves had more influence on paleoelevation reconstruction than that in other parameters".

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.
This expired link is available through the Internet Archive´s Wayback Machine.

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