Home / Palaeoclimate / Leaf Size and Shape and the Reconstruction of Past Climates

L.S.R. Alves et al. (2005): Paleobotany and Paleoclimatology Part I: Growth Rings in Fossil Woods and Paleoclimates. PDF file; See also starting with PDF-page 16:
Part II: Leaf Assemblages (Taphonomy, Paleoclimatology and Paleogeography). In: pp 179–202, Koutsoukos, Eduardo A.M. (ed.) Applied Stratigraphy. Series: Topics in Geobiology, Vol. 23.
See also here (Google books).

American Society of Plant Biologists, The Plant Cell Online: Leaf Development 1 and Leaf Development 2 (Cell proliferation and differentiation). Lecture notes, PDF files.
Snapshots provided by the Internet Archive´s Wayback Machine.
For PowerPoint Slide Presentations see here.

! Nan Crystal Arens, C. Strömberg and A. Thompson, Department of Integrative Biology, and Paleobotany Section, Museum of Paleontology (UCMP), University of California at Berkeley: Virtual Paleobotany, Lab XI. Interpreting ancient climate from fossil assemblage (e.g. climate-leaf analysis multivariate program: CLAMP; nearest living relative or the coexistence model; leaf margin analysis).

K.L. Bacon et al. (2013): Increased Atmospheric SO₂ Detected from Changes in Leaf Physiognomy across the Triassic-Jurassic Boundary Interval of East Greenland. In PDF, Plos One, 8.

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.

! Museum of Paleontology, University of California, Berkeley: The Cleared Leaf Collection. An image gallery of modern leaves that have been bleached and stained to make their venation patterns more visible. Leaf shape, venation, and features of the margin, base and apex constitute important taxonomic and physiognomic characters. See also here.

D.J. Beerling et al. (2001): Evolution of leaf-form in land plants linked to atmospheric CO₂ decline in the Late Palaeozoic era. PDF file, Nature, 410.

D.J. Beerling (1998): The future as the key to the past for palaeobotany? Abstract, Trends in Ecology & Evolution.
This expired link is available through the Internet Archive´s Wayback Machine.

! Museum of Paleontology, University of California, Berkeley:
The Cleared Leaf Collection. Excellent!
An image gallery of modern leaves that have been bleached and stained to make their venation patterns more visible. Leaf shape, venation, and features of the margin, base and apex constitute important taxonomic and physiognomic characters.
You can search the collection from the
Paleontology Collections Photos page or
the collection at the Modern Cleared Leaf Photos page.
Don't miss the helpful
! Manual of Leaf Architecture. In PDF.

E. Biffin et al. (2013): Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation. In PDF, Proc. R. Soc. B, 279: 341-348.

B. Blonder and B.J. Enquist (2014): Inferring climate from angiosperm leaf venation networks. In PDF, New Phytologist.

! B. Blonder et al. (2012): The leaf-area shrinkage effect can bias paleoclimate and ecology research. Free access, American Journal of Botany, 99: 1756-1763.

B. Blonder et al. (2012): X-ray imaging of leaf venation networks. In PDF, New Phytologist.

! B. Blonder et al. (2011): Venation networks and the origin of the leaf economics spectrum. In PDF, Ecology Letters, 14: 91-100. See also here.

Botany.Com, the Encyclopedia of Plants:
Leaf shapes.
Still available via Internet Archive Wayback Machine.

Department of Geological Sciences, University of Colorado, Boulder, CO: Web-based instruction. Annotated links to information on using the web to teach. Go to: CzPaleobotany. Go to: Cenozoic Elevation of the Rocky Mountains, Paleobotanical Methods. About fossil classification (nearest living relative, physiognomy and CLAMP) and climate and elevation analysis.
These expired links are now available through the Internet Archive´s Wayback Machine

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 M.A. Zwieniecki (2012): Leaf fossil record suggests limited influence of atmospheric CO₂ on terrestrial productivity prior to angiosperm evolution. Free access, PNAS, 109: 10403–10408.

C. Kevin Boyce et al. (2010): Angiosperms Helped Put the Rain in the Rainforests: The Impact of Plant Physiological Evolution on Tropical Biodiversity. PDF file, Annals of the Missouri Botanical Garden, 97: 527-540.
Provided by the Internet Archive´s Wayback Machine.

! C. Kevin Boyce et al. (2009). Angiosperm leaf vein evolution was physiologically and environmentally transformative. PDF file, Proceedings of the Royal Society B, 276: 1771-1776. See also here (abstract).

! C.K. Boyce (2008): The fossil record of plant physiology and development: what leaves can tell us. In PDF, Paleontological Society Papers, 14: 133–146.

! C.K. Boyce (2008): Seeing the forest with the leaves-clues to canopy placement from leaf fossil size and venation characteristics. In PDF, Geobiology, 7: 192-199.
See also here.

J. Bres et al. (2021): The Cretaceous physiological adaptation of angiosperms to a declining _pCO₂: a modeling approach emulating paleo-traits. Free access, Biogeosciences, 18: 5729–5750.
"... we show that protoangiosperm physiology does not allow vegetation to grow under low _pCO₂
[...] confirms the hypothesis of a likely evolution of angiosperms from a state of low leaf hydraulic and photosynthetic capacities at high _pCO₂ to a state of high leaf hydraulic and photosynthetic capacities linked to leaves with more and more veins together ..."

! 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).

Tim J. Brodribb et al. (2010): Viewing leaf structure and evolution from a hydraulic perspective. PDF file, Functional Plant Biology, 37: 488-498.

T.J. Brodribb and T.S. Feild (2010): Leaf hydraulic evolution led a surge in leaf photosynthetic capacity during early angiosperm diversification. In PDF, Ecology Letters, 13: 175-183. See also here.
"... Our data suggest that early terrestrial angiosperms produced leaves with low photosynthetic rates, but that subsequent angiosperm success is linked to a surge in photosynthetic capacity during their early diversification".

W.K. Buechler (2014): Variability of venation patterns in extant genus: Implications for fossil taxonomy. PaleoBios, 30: 89-104.

R.J. Burnham et al. (2001): Habitat-related error in estimating temperatures from leaf margins in a humid tropical forest. PDF file, American Journal of Botany, 88: 1096-1102.

R.J. Burnham (1994): Patterns in tropical leaf litter and implications for angiosperm paleobotany. In PDF, Review of Palaeobotany and Palynology.

M.J. Butrim et al. (2022): No Consistent Shift in Leaf Dry Mass per Area Across the Cretaceous—Paleogene Boundary. Front. Plant Sci., 13:894690. doi: 10.3389/fpls.2022.894690. See also here.

M. Coiro et al. (2024): Parallel evolution of angiosperm-like venation in Peltaspermales: a reinvestigation of Furcula. Open access, New Phytologist, doi: 10.1111/nph.19726.
"... Although a hierarchical-reticulate venation also occurs in some groups of extinct seed plants, it is unclear whether these are stem relatives of angiosperms
[...] We further suggest that the evolution of hierarchical venation systems in the early Permian, the Late Triassic, and the Early Cretaceous represent ‘natural experiments’ that might help resolve the selective pressures enabling this trait to evolve ..."

! 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.

D.L. Contreras (2018): A workflow and protocol describing the field to digitization process for new project-based fossil leaf collections. Open access, Applications in Plant Sciences, 6: e1025.

! J.S. Cope et al. (2012): Plant species identification using digital morphometrics: A review. In PDF, Expert Systems with Applications, 39: 7562-7573.
See also here.
"... We review the main computational, morphometric and image processing methods
[...] We discuss the measurement of leaf outlines, flower shape, vein structures and leaf textures, and describe a wide range of analytical methods in use.

H.J. de Boer et al. (2012): A critical transition in leaf evolution facilitated the Cretaceous angiosperm revolution. In PDF, Nature Communications, 3. See also here.

! Denver Museum of Nature and Science, Denver, Colorado: DMNS Paleobotany Collection. This website contains over 1000 images of fossil plants spanning the late Cretaceous through early Eocene from the Western Interior of North America. Go to: Identification Flow Chart, or start with Morphotype a Flora. A guide to morphotyping (or binning) a fossil flora step-by-step.

! Digiphyll (State Museum of Natural History Stuttgart).
Digiphyll is designed as an educational portal to provide effective assistance in identifying fossil plant material. Excellent! Please note:
Manual: How to use this portal (in PDF).
Glossary: Leaf morphology (in PDF).
! Worth to check out: The fact sheets (PDF files, 36 taxa). Click the button "Downloads".

! 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 ..."

David L. Dilcher, Paleobotany Laboratory, Florida Museum of Natural History, University of Florida, Gainesville, FL: Dilcher's Swamp/Woods Leaf Images.

V.M. Dörken, FB Biologie, Universität Konstanz: Morphologie und Anatomie des Blattes. PDF file, in German.
Still available via Internet Archive Wayback Machine.

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.

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.

B. Ellis and K.R. Johnson (2013): Comparison of leaf samples from mapped tropical and temperate forests: Implications for interpretations of the diversity of fossil assemblages. Abstract, Palaios.

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.

! Constantin von Ettingshausen (1858): Die Blattskelete der Apetalen, eine Vorarbeit zur Interpretation der fossilen Pflanzenreste (in German). Provided by Google books.

H.J. Falcon-Lang and D.J. Cantrill (2001): Leaf phenology of some mid-Cretaceous polar forests, Alexander Island, Antarctica. In PDF, Geological Magazine.

T.S. Feild et al. (2011): Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution. In PDF, PNAS, 108: 8363-8366.

Ian J. Glasspool et al.: Foliar physiognomy in Cathaysian gigantopterids and the potential to track Palaeozoic climates using an extinct plant group. Palaeogeography, Palaeoclimatology, Palaeoecology, 205: 69-110; 2004.

W.A. Green et al. (2015): Reading the leaves: A comparison of leaf rank and automated areole measurement for quantifying aspects of leaf venation. In PDF.

D.R. Greenwood (2007): Fossil angiosperm leaves and climate: from Wolfe and Dilcher to Burnham and Wilf. In PDF, Courier Forsch.-Senckenberg, 258.
The link is to a version archived by the Internet Archive´s Wayback Machine.

David R. Greenwood, Environmental Science, Brandon University, Canada: Commentary - Leaf form and the reconstruction of past climates (Commentary on Traiser et al. 2005). Abstract, New Phytologist, 166, 355-357; 2005.

G.W. Grimm and A.J. Potts (2015): Fallacies and fantasies: the theoretical underpinnings of the Coexistence Approach for palaeoclimate reconstruction. In PDF, Clim. Past Discuss., 11: 5727-5754.

G.W. Grimm et al. (2015): Fables and foibles: a critical analysis of the Palaeoflora database and the Coexistence approach for palaeoclimate reconstruction. In PDF.

S.G. Hao and J.Z. Xue (2013): Earliest record of megaphylls and leafy structures, and their initial diversification. In PDF, Chin. Sci. Bull., 58: 2784-2793.

! E.R. Hagen et al. (2019): No Large Bias within Species between the Reconstructed Areas of Complete and Fragmented Fossil Leaves. Abstract, Palaios, 34: 43-48. See also here (in PDF).
"... that the underrepresentation of large leaves, as captured by our study design, is probably not critical for most fossil applications. Comparing directly the reconstructed areas of complete and fragmented leaves appears reasonable, thus expanding the usefulness of fossil leaf fragments. ..."

! L.J. Hickey&xnbsp;(1973): Classification of the architecture of dicotyledonous leaves. In PDF, American journal of botany, 60: 17-33.
See also here.
! Note figure 1-40: Leaf orientation features: Orientation and form of whole leaf, shape of apex and base, gland position, and marginal configuration.
! Figure 41-62: Types of venation.
! Figure 63-87: Orders of venation and vein configuration.
! Figure 88-107: Ultimate venation and areolation.

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).

M. Hrabovský (2021): Leaf evolution and classification. 3. Gymnospermopsida. In PDF, Acta Botanica Universitatis Comenianae, 57.
! Many black and white contour drawings.

M. Hrabovský (2020): Leaf evolution and classification. 2. Polypodiopsida. In PDF, Acta Botanica Universitatis Comenianae, 56.
! Many black and white contour drawings.

M. Hrabovský (2020): Leaf evolution and classification. 1. Lycopodiopsida. In PDF, Acta Botanica Universitatis Comenianae, 55.
See also here.
! Many black and white contour drawings.

P.M. Huff et al. (2003): Digital future for paleoclimate estimation from fossil leaves? Preliminary results. PDF file, Palaios, 18: 266-274.

! M.E. James et al. (2023): Replicated Evolution in Plants. Open access, Annual Review of Plant Biology, 74: 697-725.
"...Similar traits and functions commonly evolve in nature. Here, we explore patterns of replicated evolution across the plant kingdom and discuss the processes responsible for such patterns.
[...] The term replicated evolution can be used to encompass both convergence and parallelism ..."

! G.J. Jordan (2011): A critical framework for the assessment of biological palaeoproxies: predicting past climate and levels of atmospheric CO2 from fossil leaves. In PDF, New Phytologist.

N.A. Jud and L.J. Hickey (2013): Potomacapnos apeleutheron gen. et sp. nov., a new Early Cretaceous angiosperm from the Potomac Group and its implications for the evolution of eudicot leaf architecture. In PDF, Am. J. Bot., see also here. ^

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.

W. Konrad et al. (2021): Leaf temperature and its dependence on atmospheric CO₂ and leaf size. Open access, Geological Journal, 56.

E.A. Kowalski and D.L. Dilcher (2002): Warmer paleotemperatures for terrestrial ecosystems. In PDF, PNAS, 100: 167-170.

Jonathan Krieger, Robert Guralnick, Kirk Johnson & Dena Smith: Predicting climate using empirically determined continuous measures of leaf shape. Abstract, Botany 2004, The Botanical Society of America. See also here.

! M. Li et al. (2017): Persistent homology demarcates a leaf morphospace. In PDF, bioRxiv. See also here.

S.A. Little et al. (2014): Reinvestigation of Leaf Rank, an Underappreciated Component of Leo Hickey´s Legacy. In PDF.

! S.A. Little et al.(2010): Paleotemperature Proxies from Leaf Fossils Reinterpreted in Light of Evolutionary History. In PDF, PLoS ONE, 5: e15161. See also here.

Z. Lv et al. (2023): Overview of molecular mechanisms of plant leaf development: a systematic review. Free access, Frontiers in Plant Science, 14.

! H. Ma et al. (2023): The global biogeography of tree leaf form and habit. Open access, Nature Plants, https://doi.org/10.1038/s41477-023-01543-5.

L. Mander and H.T.P. Williams (2024): The robustness of some Carboniferous fossil leaf venation networks to simulated damage. Open access, R. Soc. Open Sci. 11: 240086. https://doi.org/10.1098/rsos.240086.
"... We attacked fossil venation networks with simulated damage to individual vein segments and leaf blades. For both types of attack, branched venation networks are the least robust to damage, with greater robustness shown by the net-like reticulate networks
[...] A living angiosperm Betula alba was the most robust in our analysis ..."

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.

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.

M.L. McKee et al. (2019): Experimental evidence for species-dependent responses in leaf shape to temperature: Implications for paleoclimate inference. Open access, PLoS ONE, 14: e0218884.

! MORPYHLL - database for acquisition of ecophysiologically relevant morphometric data of fossil leaves.
See also here. Please note:
C. Traiser et al. (2018): MORPHYLL: A a database of fossil leaves and their morphological traits. Palaeontologia Electronica, 21.1.1T: 1-17. Available in PDF.

! V. Mosbrugger and T. Utescher (1997): The coexistence approach -- a method for quantitative reconstructions of Tertiary terrestrial palaeoclimate data using plant fossils. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 134: 61-86.
See also here.

! C. Müller et al. (2023): An integrated leaf trait analysis of two Paleogene leaf floras. In PDF, PeerJ 11: e15140 https://doi.org/10.7717/peerj.15140.
See also here.
Note figure 1: Schematic overview of the datasets used and their selection process.
Figure 6: Herbivory metrics compared between Seifhennersdorf and Suletice-Berand regarding whole assemblages and fossil-species phenology.
"... This study presents the Integrated Leaf Trait Analysis (ILTA), a workflow for the combined application of methodologies in leaf trait and insect herbivory analyses on fossil dicot leaf assemblages ..."

A.B. Nicotra et al. (2011): The evolution and functional significance of leaf shape in the angiosperms. In PDF, Functional Plant Biology, 38: 535-552. See also here.

K.J. Niklas (1999): A mechanical perspective on foliage leaf form and function. In PDF, New Phytologist.

Ülo Niinemets et al. (2007): Do we Underestimate the Importance of Leaf Size in Plant Economics? Disproportional Scaling of Support Costs Within the Spectrum of Leaf Physiognomy. PDF file, Ann. Bot., 100: 283-303.

D.A. Oliva et al. (2022): First record of plant macrofossil from the Boa Vista Formation, Takutu Basin, Roraima State, Brazil. In PDF, Revista Brasileira de Paleontologia, 25: 303–321.
See also here.
"... X-ray diffractometry (XRD) and Laser induced-breakdown spectroscopy (LIBS) analysis were performed ..."

Sofia Oliver (2010): Digital leaf physiognomy: correlating leaf size and shape to climate in the Fox Hills, Fort Union, and Hanna Basin Formations. PDF file; Thesis, Wesleyan University.

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.

! D.J. Peppe (2018): Reconstructing paleoclimate and paleoecology using fossil leaves. Abstract, in: Croft D., Su D., Simpson S. (eds) Methods in Paleoecology. Vertebrate Paleobiology and Paleoanthropology. Springer. See also here (in PDF).

! D.J. Peppe et al. (2011): Sensitivity of leaf size and shape to climate: global patterns and paleoclimatic applications. Free access, New Phytologist, 190: 724-739.

! 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.

! Christian Pott and Michael Krings (2010): Gymnosperm Foliage from the Upper Triassic of Lunz, Lower Austria: an annotated check list and identifiation key. PDF file, Geo.Alp, 7: 19-38.

M.P. Pound et al. (2017): Deep Machine Learning provides state-of-the-art performance in image-based plant phenotyping. GigaScience. See also here (in PDF).

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

! C.A. Price et al. (2011): Scaling and structure of dicotyledonous leaf venation networks. In PDF, Ecology Letters.
See also here.

Charles A. Price et al. (2011): Leaf Extraction and Analysis Framework Graphical User Interface: Segmenting and Analyzing the Structure of Leaf Veins and Areoles. Plant Physiol., 155: 236-245.

Radboud University, Nijmegen, The Netherlands: Virtual Classroom Biology. This website is an educational site, especially meant for secondary school students who like to have a first glance on teaching items of the Bio-science programs. One can find custom-made teaching material for courses from the biology training. Navigate from here. See especially:
The Microworld of Leaves.

! A.E. Radford, W.C. Dickison, J.R. Massey, & C.R. Bell (Harper and Row, New York): Vascular Plant Systematics. This book was written as a reference text for basic courses in taxonomy and as a source book of information, procedures and references for ecosystematics, biosystematics, phylosystematics and chemosystematics. It includes (1) an essentially synoptical treatment of the evidence, principles, and concepts considered fundamental to vascular plant taxonomic studies and research;
(2) organized laboratory and field exercises and problems basic to systematics;
(3) useable and useful techniques;
(4) summaries of terminology pertinent to taxonomy;
(5) relevant bibliographies and indices; and (6) information on systematic facilities.
Searching images you may navigate from here. See also:
! Section A. Structure and Specialized Characters: V. Leaves, or Section B: General Characters and Character States, A. Location or Environmental Position. Classification based on position of organs or parts in their surrounding environment.

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

! G. Rossetto-Harris et al. (2022): Rapid character scoring and tabulation of large leaf-image libraries using Adobe Bridge. Open access, Appl. Plant Sci., 10: e11500.
Note figure 1: Flowchart illustrating the workflow to annotate large image libraries.
"... Our approach is intuitive and acts as a digital mimic and complement to the experience of sorting and analyzing specimens in-person. Keywords can be easily customized for other data types that require visual sorting using image libraries ..."

A. Roth-Nebelsick and C. Traiser (2024): Diversity of leaf architecture and its relationships with climate in extant and fossil plants. In PDF. Palaeogeography, Palaeoclimatology, Palaeoecology, 634.
See also here.
"... the diversity of functional leaf architecture and its association with climate is studied for extant woody dicot species
[...] results of this study indicate that diversity of leaf architecture may be a useful source of information for palaeoecology and palaeoclimate ..."

A. Roth-Nebelsick et al. (2021): Taxon-specific variability of leaf traits in three long-ranging fossil-species of the Paleogene and Neogene: Responses to climate. In PDF, Palaeontologia Electronica, 24: a04. https://doi.org/10.26879/1114.

Anita 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. (2012): Roles of climate and functional traits in controlling toothed vs. untoothed leaf margins. In PDF, American Journal of Botany, 99: 915-922.

! D.L. Royer (2012): Climate reconstruction from leaf size and shape: New developments and challenges. PDF file, in: Reconstructing Earth´s Deep-Time Climate - The State of the Art in 2012, Paleontological Society Short Course, The Paleontological Society Papers, Volume 18, Linda C. Ivany and Brian T. Huber (eds.), pp. 195-212.

D. Royer et al. (2010): Leaf economic traits from fossils support a weedy habit for early angiosperms. Free access, American Journal of Botany, 97: 438-445.

Dana L. Royer et al. (2009): Phenotypic Plasticity of Leaf Shape along a Temperature Gradient in Acer rubrum. PDF file.

D.L. Royer et al. (2009): Ecology of leaf teeth: A multi-site analysis from an Australian subtropical rainforest. PDF file, American Journal of Botany, 96: 738–750.

D.L. Royer et al. (2012): ! Roles of climate and functional traits in controlling toothed vs. untoothed leaf margins. In PDF, American Journal of Botany, 99: 915-922.

! D.L. Royer et al. (2005): Correlations of climate and plant ecology to leaf size and shape: potent.ial proxies for the fossil record. Open access, American Journal of Botany, 92: 1141-1151.

Royer et al.: DIGITAL LEAF PHYSIOGNOMY: CALIBRATION OF A NEW METHOD FOR RECONSTRUCTING CLIMATE FROM FOSSIL PLANTS. Abstract, 2004 GSA Denver Annual Meeting.

L. Sack et al. (2013): How do leaf veins influence the worldwide leaf economic spectrum? Review and synthesis. Journal of Experimental Botany, 64: 4053-4080.

! L. Sack and C. Scoffoni (2013): Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future. Free access, New Phytologist, 198: 983–1000.
Note figure 6: Evolution of terrestrial plants and their traits, including leaf vein traits against geological periods and time.

! L. Sack et al. (2012): Developmentally based scaling of leaf venation architecture explains global ecological patterns. Free access, Nature Communications, 3.
Note figure 4: The geometric scaling of vein density with leaf size expected because of leaf expansion and mechanisms for its modification.

H.L. Sanders and S.E. Wyatt (2009): Leaf Evolution and Development: Advancing Technologies, Advancing Understanding. Free access, BioScience, 59: 17-26.

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, Dissertation, Lund University.

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 ..."

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).

C.M. Sosa and J.G. Puntieri (2023): Are scale leaves essencial in temperate-cold climates? An evaluation in tree species from temperate rainforests of South America. Open access, Acta Botanica Brasilica, 37.

R.A. Spicer et al. (2009): New developments in CLAMP: Calibration using global gridded meteorological data. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 283: 91-98.

! 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. Spicer, Palaeoenvironmental Research Group, Earth Sciences Dept., The Open University, Milton Keynes: CLAMP Online. CLAMP is a method of obtaining ancient climate information from the architecture (physiognomy) of fossil leaves of woody dicot flowering plants.

! Robert A. Spicer, Earth Sciences Department, The Open University, Milton Keynes, U.K. (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. Go to: CLAMP Leaf Character State Definitions and Scoring, and Leaf Size Template.

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.

M. Tanrattana et al. (2020). Climatic evolution in Western Europe during the Cenozoic: insights from historical collections using leaf physiognomy. In PDF, Geodiversitas, 42: 151-174.
See also here and there.

! Vasilis Teodorides et al. (2011): Refining CLAMP - investigations towards improving the Climate Leaf Analysis Multivariate Program. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology.

V. Teodoridis et al. (2011): The integrated plant record vegetation analysis: internet platform and online application. In PDF, Acta Musei Nationalis Pragae, Ser. B, 67: 159-165.

A.M.F. Tomescu (2009): Megaphylls, microphylls and the evolution of leaf development. PDF file, Trends in plant science, 14.

! A. Toumoulin et al. (2020): Reconstructing leaf area from fragments: testing three methods using a fossil paleogene species. In PDF, American Journal of Botany, 107: 1786–1797. See also here.

Christopher Traiser, Tübingen University: Blattphysiognomie als Indikator für Umweltparameter: Eine Analyse rezenter und fossiler Floren (Thesis, PDF file, in German). This study investigates the relationship between physiognomic traits of leaves from European hardwood vegetation and environmental parameters in order to create a calibration dataset. The leaf data are obtained from synthetic chorologic floras, the environmental data comprise climatic and ecologic data.

C. Traiser et al. (2005): Environmental signals from leaves - a physiognomic analysis of European vegetation. Free access, New Phytologist, 166: 465-484.

D. Uhl et al. (2007): Cenozoic paleotemperatures and leaf physiognomy - A European perspective. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 248: 24-31.

! D. Uhl (2006): Fossil plants as palaeoenvironmental proxies - some remarks on selected approaches. PDF file, Acta Palaeobotanica, 46: 87-100.

! T. Utescher et al. (2014): The Coexistence Approach - Theoretical background and practical considerations of using plant fossils for climate quantification. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology. 410: 58-73.
Snapshot provided by the Internet Archive´s Wayback Machine.

! Johanna H.A. van Konijnenburg-van Cittert (2008): The Jurassic fossil plant record of the UK area. PDF file, Proceedings of the Geologists' Association 119: 59-72.
! See fig. 6: how to distinguish bennettialean leaf shapes!
Now provided by the Internet Archive´s Wayback Machine.

! Visual Plants. Based on a scientific database, the program can be used for the visual determination of plants. Now with around 22000 images, mostly with geo-referenced information. You can search using taxon names or via plant characters.
Worth checking out: Dalitz, H. and Homeier, J. (2004): Visual Plants - An image based tool for plant diversity research. Lyonia, 6: 47-59.
See also here (in German).

H. Wang et al. (2019): Plant leaf tooth feature extraction. Open access, PLoS ONE, 14: e0204714.

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.

M.C. Wiemann et al. (1998): Estimation of temperature and precipitation from morphological characters of dicotyledonous leaves. In PDF, American Journal of Botany, 85: 1796–1802. See also here.

Wikipedia, the free encyclopedia:
! Leaf.

Wikipedia, the free encyclopedia:
Blattpolymorphismus (in German).

Wikipedia, the free encyclopedia:
! Category:Glossaries of botany.
! Glossary of leaf morphology.

P. Wilf et al. (2021): An image dataset of cleared, x-rayed, and fossil leaves vetted to plant family for human and machine learning. Open access, PhytoKeys, 187: 93–128. Go to:
! Dataset (available from the Figshare Plus repository). Image collection and supporting data for: An image dataset of cleared, x-rayed, and fossil leaves vetted to plant family for human and machine learning.
See also:
From museum to laptop: Visual leaf library a new tool for identifying plants (by Matthew Carroll, March 15, 2022).
Penn State: From museum to laptop: Visual leaf library a new tool for identifying plants.

P. Wilf (2008): Fossil angiosperm leaves: paleobotany´s difficult children prove themselves. PDF file, Paleontological Society Papers, 14: 319-333.

P. Wilf et al. (1998): Using fossil leaves as paleoprecipitation indicators: an Eocene example. In PDF, Geology,26: 203-206. See also here.

! P. Wilf (1997): When are leaves good thermometers? A new case for leaf margin analysis. In PDF, Paleobiology, 23: 373-390.

! C.G. Willis et al. (2017): Old Plants, New Tricks: Phenological Research Using Herbarium Specimens. In PDF, Trends in Ecology & Evolution, 32: 531-546. See also here.
"... Herbarium specimens provide a window into the past that increases our temporal, geographic – and taxonomic vision of how phenology – and potentially plant success and ecosystem processes, have changed and will continue to be affected as the climate changes. With a thorough and growing understanding of the potential and limitations of this rich historical data source, combined with the modern tools of digitization, data sharing, and integration, researchers will increasingly be able to address critical questions about plant biology ..."

M.C. Wiemann et al. (2018): Estimation of temperature and precipitation from morphological characters of dicotyledonous leaves. Free access, American Journal of Botany, 85: 11796-1802.

Wired, Boone, IA: A Computer With a Great Eye Is About to Transform Botany (March 17, 2016).
See also here (Geological Society of America Abstracts) and there P. Wilf et al., Computer vision cracks the leaf code (PNAS).

! J.A. Wolfe and G.R. Upchurch (1987): Leaf assemblages across the Cretaceous-Tertiary boundary in the Raton Basin, New Mexico and Colorado. Free access, Proc. National Academy of Sciences USA, 84: 5096-5100.

Jack A. Wolfe (1978): A Paleobotanical Interpretation of Tertiary Climates in the Northern Hemisphere: Data from fossil plants make it possible to reconstruct Tertiary climatic changes, which may be correlated with changes in the inclination of the earth's rotational axis. In PDF, American Scientist, 66: 694-703.

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".

! J. Yang et al. (2015): Leaf form-climate relationships on the global stage: an ensemble of characters. In PDF, Global Ecology and Biogeography.

! Y.J. Zhang et al. (2015): Extending the generality of leaf economic design principles in the cycads, an ancient lineage. Free access, New Phytologist, 206: 817–829.

Y. Xu et al. (2023): How similar are the venation and cuticular characters of Glossopteris, Sagenopteris and Anthrophyopsis? In PDF, Review of Palaeobotany and Palynology, 316.
See likewise here.
Note figure 1: Geologic ranges of some representative reticulate taxa.
"... Considering the putatively close relationship of glossopterids (Glossopteris), Caytoniales (Sagenopteris) and Bennettitales (here encompassing Anthrophyopsis) resolved as members of the ‘glossophyte’ clade in some past phylogenetic studies, cuticular features suggest that these groups are not closely related. In addition, anastomosing venation, superficially similar to that of Glossopteris, Sagenopteris and Anthrophyopsis appears to have arisen independently in numerous other plant groups ..."

M.A. Zwieniecki and C.K. Boyce (2014): Evolution of a unique anatomical precision in angiosperm leaf venation lifts constraints on vascular plant ecology. In PDF, Proc. R. Soc. B, 281: 20132829. See also here.

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