Leaf Size and Shape and the Reconstruction of Past Climates
Focused on Palaeoclimate
Tree-Ring Research (Dendrochronology) in General
The Pros and Cons of Pre-Neogene Growth Rings
The Rise of Oxygen and the Global Carbon Cycle
Abscission and Tissue Separation in Fossil and Extant Plants@
Teaching Documents about Botany@
Introductions to both Fossil and Recent Plant Taxa@
Laureen Sally da Rosa Alves and Margot Guerra-Sommer (2007):
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: Koutsoukos, Eduardo A.M. (ed.) Applied Stratigraphy. Series: Topics in Geobiology, Vol. 23.
See also here (in PDF) and there (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.
! 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):
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.
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 In PDF, 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.
Botany.Com, the Encyclopedia of Plants: Leaf shapes. Snapshot taken by the Internet Archive´s 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.
C.K. Boyce and M.A. Zwieniecki (2012): Leaf fossil record suggests limited influence of atmospheric CO2 on terrestrial productivity prior to angiosperm evolution. In PDF, PNAS, 109.
C. Kevin Boyce et al. (2010):
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.
! 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):
hydraulic evolution led a surge in leaf
photosynthetic capacity during early angiosperm
diversification. In PDF, Ecology Letters, 13: 175-183.
"... 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.
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.
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.
! D.L. Dilcher (1974): Approaches to the identification of angiosperm leaf remains. In PDF, The Botanical Review, 40: 1–157. See also here.
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. In German.
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. First part of a reference book of the Tertiary flora of Saxony. See also here (abstract).
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):
of Leaf Architecture. Book announcement.
! See also here.
! 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.
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.
P.M. Huff et al. (2003): Digital future for paleoclimate estimation from fossil leaves? Preliminary results. PDF file, Palaios, 18: 266-274.
! 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.
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.
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.
! 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.
K.J. Niklas (1999): A mechanical perspective on foliage leaf form and function. In PDF, New Phytologist.
! 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.
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.
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):
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 et al. (2011): Sensitivity of leaf size and shape to climate: global patterns and paleoclimatic applications. In PDF, New Phytologist, 190: 724-739. See also here (abstract).
! 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.
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.
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.
Jennifer Read and Alexia Stokes (2006): Plant biomechanics in an ecological context. PDF file, American Journal of Botany, 93: 1546-1565.
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. PDF file, American Journal of Botany, 97: 438-445. See also here.
Dana L. Royer et al. (2009): Phenotypic Plasticity of Leaf Shape along a Temperature Gradient in Acer rubrum. PDF file.
Dana 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.
Royer DL, Wilf P, Janesko DA, Kowalski EA, Dilcher DL. (in press): Correlations of climate and plant ecology to leaf size and shape: potential proxies for the fossil record. PDF file, Amer J Bot (in press).
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. In PDF, New Phytologist, 198: 983–1000.
L. Sack et al. (2012): Developmentally based scaling of leaf venation architecture explains global ecological patterns. In PDF.
H.L. Sanders and S.E. Wyatt (2009): Leaf Evolution and Development: Advancing Technologies, Advancing Understanding. PDF file, 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.
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 et al. (2009): New developments in CLAMP: Calibration using global gridded meteorological data. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 283: 91-98.
! 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.
! 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. 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.! C. Traiser et al. (2018): MORPHYLL: A database of fossil leaves and their morphological traits. Palaeontologia Electronica. See also here (in PDF).
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.
Traiser, C, Klotz, S., Uhl, D., & Mosbrugger, V. (2005). Environmental signals from leaves - a physiognomic analysis of European vegetation. PDF file, 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.
! 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.
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).
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. American Journal of Botany, 85: 1796–1802.
Wikipedia, the free encyclopedia:
the free encyclopedia:
Blattpolymorphismus (in German).
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):
Plants, New Tricks: Phenological Research Using Herbarium Specimens. In PDF,
Trends in Ecology & Evolution, 32: 531-546.
"... 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 ..."
Wired, Boone, IA:
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).
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):
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. In PDF, New Phytologist.
M.A. Zwieniecki and C.K. Boyce (2014):
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
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