Links for Palaeobotanists

Home / Introductions to both Fossil and Recent Plant Taxa / Coniferophyta

Cyanobacteria and Stromatolites
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Picture Search@
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! J. Anderson et al. (2007): Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology. In PDF, Strelitzia, 20, 279 p. See also here (abstract).

A. Andruchow-Colombo et al. (2023): In search of lost time: tracing the fossil diversity of Podocarpaceae through the ages. In PDF, Botanical Journal of the Linnean Society.
See also here.

A. Andruchow-Colombo et al. (2022): New genus of Cupressaceae from the Upper Cretaceous of Patagonia (Argentina) fills a gap in the evolution of the ovuliferous complex in the family. In PDF, Journal of Systematics and Evolution.
See also here.

A. Andruchow-Colombo et al. (2019): Oldest record of the scale-leaved clade of Podocarpaceae, early Paleocene of Patagonia, Argentina. In PDF, Alcheringa, 43. See also here.

A. Andruchow-Colombo et al. (2019): A South American fossil relative of Phyllocladus: Huncocladus laubenfelsii gen. et sp. nov.(Podocarpaceae), from the early Eocene of Laguna del Hunco, Patagonia, Argentina. Australian Systematic Botany, 32: 290–309. See also here (in PDF).

! 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. Go to: Ginkgo, Cordaites and the Conifers.

Wayne P. Armstrong, Pacific Horticulture: The Araucaria Family: Past & Present. Please take notice the diorama of an araucariad forest from 200 million years ago (Diorama on display at the Rainbow Forest Museum, Petrified Forest National Park).

B.J. Axsmith and S.R. Ash (2006): Two rare fossil cones from the Upper Triassic Chinle Formation in Petrified Forest National Park, Arizona, and New Mexico. In PDF, Museum of Northern Arizona Bulletin, 62.

B.J. Axsmith et al. (1998): A New Fossil Conifer from the Triassic of North America: Implications for Models of Ovulate Cone Scale Evolution. PDF file, International Journal of Plant Sciences.
See also here.

B.J. Axsmith et al. (1998): Anatomically preserved leaves of the conifer Notophytum krauselii (Podocarpaceae) from the Triassic of Antarctica. In PDF, American Journal of Botany, 85: 704–713. See also here and there.

Brian J. Axsmith and Thomas N. Taylor (1997): The Triassic conifer seed cone Glyptolepis. PDF file.
Now recovered from the Internet Archive´s Wayback Machine.

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

M.E.P. Batista et al. (2020): A New Species of Brachyphyllum from the Crato Formation (Lower Cretaceous), Araripe Basin, Brazil. In PDF, Ameghiniana, 57: 519-533. See also here.

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

N.N.A. Bayam et al. (2018): Further contributions to the early Miocene forest vegetation of the Galatian Volcanic Province, Turkey. Free access, Palaeontologia Electronica.

N.V. Bazhenova et al.(2022): Mummified Seed Cones of Pinus prehwangshanensis sp. nov. (Subgenus Pinus, Pinaceae) From the Upper Pleistocene of Guangdong, South China: Taxonomical Significance and Implication for Phytogeography and Ecology. Free access, Front. Ecol. Evol., 10: 900687. doi: 10.3389/fevo.2022.900687

J.A. Bergene (2012): Dordrechtites arcanus, an anatomically preserved gymnospermous reproductive structure from the Middle Triassic of Antarctica. In PDF, thesis, University of Kansas.

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.

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

J. Bodnar and I.H. Escapa (2016): Towards a whole plant reconstruction for Austrohamia (Cupressaceae): New fossil wood from the Lower Jurassic of Argentina. Abstract, Review of Palaeobotany and Palynology, 234: 186-197. See also here (in PDF).
Note Figure 2: The vegetation of the Cerro Bayo landscape (Early Jurassic, Patagonia), consisting mainly of Austrohamia minuta. In the understorey dipteridaceous, osmundaceous and marattiaceous ferns.

! B. Bomfleur et al. (2013): Whole-Plant Concept and Environment Reconstruction of a Telemachus Conifer (Voltziales) from the Triassic of Antarctica. In PDF, Int. J. Plant Sci., 174: 425–444. See also here (abstract).
Note fig. 8 (PDF page 16): Reconstructions of various organs of the Triassic conifer Telemachus.

B. Bomfleur et al. (2011): The possible pollen cone of the Late Triassic conifer Heidiphyllum/Telemachus (Voltziales) from Antarctica. Abstract.

! L.M. Bowe et al. (2000): Phylogeny of seed plants based on all three genomic compartments: Extant gymnosperms are monophyletic and Gnetales' closest relatives are conifers. In PDF, PNAS, 97: 4092–4097. See also here.

G.E. Burrows et al. (2007): An Anatomical Assessment of Branch Abscission and Branch-base Hydraulic Architecture in the Endangered Wollemia nobilis. PDF file, Annals of Botany, 99: 609-623. See also here (abstract).

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

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

Mick Casper, The Lovett Pinetum Charitable Foundation, Conifer Paleobotany Basics. Snapshot taken by the Internet Archive´s Wayback Machine. See also: "Living" Paleobotany.

The Catalogue of Life.
This is the most comprehensive and authoritative global index of species currently available, consisting of a single integrated species checklist and their taxonomic hierarchy. With essential information on the names, relationships and distributions of over 1.6 million species. See especially:
Phylum Pinophyta.

Ralph W. Chaney: A Revision of Fossil Sequoia and Taxodium in Western North America Based on the Recent Discovery of Metasequoia Transactions of the American Philosophical Society, New Series, Vol. 40, No. 3 (1950), pp. 171-263. Accessible via Google Book Search.

M. Cheek (2016): Kew´s successful year of discoveries. Scroll down to "Fossil discovery".
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

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

! F.L. Condamine et al. (2020): The rise of angiosperms pushed conifers to decline during global cooling. Free access, Proceedings of the National Academy of Sciences, 117: 28867–28875.
Note figure 1: An overview of hypothetical determinants of conifer diversification over time.
Figure 2: Global diversification of conifers inferred from a molecular phylogeny and the fossil record.
Figure 3: Drivers of conifer diversification dynamics.

! D. Contreras et al. (2019): Reconstructing the Early Evolution of the Cupressaceae: A Whole-Plant Description of a New Austrohamia Species from the Cañadón Asfalto Formation (Early Jurassic), Argentina. Int. J. Plant Sci., 180: 834–868. See also here (in PDF).

! D.L. Contreras et al. (2017): Evolution of dispersal strategies in conifers: functional divergence and convergence in the morphology of diaspores. In PDF, Perspectives in Plant Ecology, Evolution and Systematics, 24: 93-117.
See also here.
"... Conifers diversified in their dispersal strategies through seed coat modifications or by the incorporation of various parts of the seed cone into the diaspore, with the modern conifer families independently evolving their characteristic diaspore compositions. Almost all functional morphotypes were present prior to the Cenozoic in at least one lineage, ..."

R. Daber and D. Müller-Doblies (2002): Männliche Zapfen des Urweltmammutbaums Metasequoia glyptostroboides (Cupressaceae)–Lebendbeobachtungen aus Berlin. PDF file, in German. Der Palmengarten.
See also here.

! C.C. Davis and H. Schaefer (2011): Plant Evolution: Pulses of Extinction and Speciation in Gymnosperm Diversity. See also here (abstract).

Owen Kent Davis, Department of Geosciences, University of Arizona Tucson: . Lecture notes. Go to: CONIFER NEEDLE EXTERNAL ANATOMY.
These expired links are now available through the Internet Archive´s Wayback Machine.

Owen Kent Davis, Department of Geosciences, University of Arizona Tucson:
Still available through the Internet Archive´s Wayback Machine.

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

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

W.N. Ding et al. (2018): A new fossil species of Cryptomeria (Cupressaceae) from the Rupelian of the Lühe Basin, Yunnan, East Asia: Implications for palaeobiogeography and palaeoecology. Abstract, Review of Palaeobotany and Palynology, 248: 41-51.
Also of interest in this context:
Pflanzliche Botschaften aus der Urzeit (by Tamara Worzewski, November 08, 2022,, in German).

dmoz, the Open Directory Project:
Science: Biology: Flora and Fauna: Plantae:
Coniferophyta. See also:
Earth Sciences: Paleontology: Paleobotany: Taxa.
These expired links are now available through the Internet Archive´s Wayback Machine.

M. Dolezych and L. Reinhardt (2020): First evidence for the conifer Pinus, as Pinuxylon selmeierianum sp. nov., during the Paleogene on Wootton Peninsula, northern Ellesmere Island, Nunavut, Canada. As well as here. In PDF, Canadian Journal of Earth Sciences, 57: 25–39.
See also here.
Note fig. 2: Type locality and holotype trunk Pinuxylon selmeierianum.

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

V.M. Dörken and H. Nimsch (2018): A monograph of leaf characters in the genus Abies (Abietoideae, Pinaceae). In PDF.

V.M. Dörken and P.J. Rudall (2018): Understanding the cone scale in Cupressaceae: insights from seed-cone teratology in Glyptostrobus pensilis. Open access, PeerJ.
Note figure 6: Schematic drawings of a single bract/scale complex in different taxodiaceous Cupressaceae.

V. Dörken (2008) Saisonalität und Langtrieb-/Kurztrieb-Differenzierung bei Gymnospermen: abgeleitet oder ursprünglich? PDF file, in German.
Seasonality and long shoot/short shoot syndrome in gymnosperms: derived or primitive? Doctoral thesis, Lehrstuhls für Evolution und Biodiversität der Pflanzen, Ruhr-Universität Bochum, Germany.
A version archived by the Internet Archive´s Wayback Machine.

A.B. Doweld (2022): (2889) Proposal to conserve the name Podozamites against Preissleria (fossil Pinophyta: Podozamitales). Free access, Taxon, 71: 484–485.
"... The fossil-generic name Podozamites, so widely used in modern systematic palaeobotany, should not be rejected for purely nomenclatural reasons. In order to stabilize palaeobotanical nomenclature in current use by legitimizing the use of Podozamites, it is formally proposed to conserve Podozamites against the “nomen oblitum”, Preissleria. ..."

! A.B. Doweld (2018): Proposal to conserve the name Glyptolepis keuperiana with a conserved type (fossil Gymnospermae: Voltziopsida). Free access, Taxon, 67.

! Christopher J. Earle (server space has been provided by the Department of Botany, Rheinische Friedrich-Wilhelms-Universität Bonn, Germany): The Gymnosperm Database. Currently the database provides basic information for all species and higher-ranked taxa of the gymnosperms, i.e., conifers, cycads, and their allies. You may navigate from the species list, alphabetized by binomial.

I. Escapa and A. Leslie (2017): A new Cheirolepidiaceae (Coniferales) from the Early Jurassic of Patagonia (Argentina): Reconciling the records of impression and permineralized fossils. Am. J. Bot., 104: 322-334. See also here (abstract).

I.H. Escapa et al. (2016): A new species of Athrotaxites (Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA. In PDF, Botany, 94: 831–845.

! I.H. Escapa and S. Catalano (2013): Phylogenetic Analysis of Araucariaceae: Integrating Molecules, Morphology, and Fossils. In PDF, International Journal of Plant Sciences. See also here.
"... Monophyletic Araucariaceae is the sister group of Podocarpaceae, forming the order Araucariales. Monophyly of Araucaria and Agathis is also strongly supported by the data. The results of both molecular and combined analyses indicate that Wollemia and Agathis form a clade (=agathioid clade) sister to Araucaria ..."

I.H. Escapa et al. (2013): Pararaucaria delfueyoi sp. nov. from the Late Jurassic Cañadón Calcáreo Formation, Chubut, Argentina: insights into the evolution of the Cheirolepidiaceae. In PDF, Int. J. Plant Sci., 174: 458-470.
The link is to a version archived by the Internet Archive´s Wayback Machine.
See also here.

I.H. Escapa et al. (2011): Seed cone anatomy of Cheirolepidiaceae (Coniferales): Reinterpreting Pararaucaria patagonica Wieland. Free access, Am. J. Bot., 99: 1058-1068.

Ignacio H. Escapa et al. (2010): Evolution and relationships of the conifer seed cone Telemachus: Evidence from the Triassic of Antarctica. PDF file, Int. J. Plant Sci., 171: 560-573.
See fig. 6: Hypothetical reconstructions of Telemachus elongatus and Telemachus antarcticus ovulate cones.

P. Falaschi et al. (2011): Growth architecture and silhouette of Jurassic conifers from La Matilde Formation, Patagonia, Argentina. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 302: 122-141.

H.J. Falcon-Lang et. al. (2016): The oldest Pinus and its preservation by fire. Abstract, Geology, 44: 303-306. See also here (in PDF).

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

H.J. Falcon-Lang et al. (2014): Coniferopsid tree trunks preserved in sabkha facies in the Permian (Sakmarian) Community Pit Formation in south-central New Mexico, U.S.A.: Systematics and palaeoecology. Abstract.

Howard J. Falcon-Lang et al. (2011): Pennsylvanian coniferopsid forests in sabkha facies reveal the nature of seasonal tropical biome. Abstract, Geology, 39: 371-374.

! A. Farjon (2018): The Kew Review: Conifers of the World. Open access, Kew Bulletin, 73: 8.

! A. Farjon (2008): A natural history of conifers. Google books, see also here.

Z. Feng et al. (2018): A conifer-dominated Early Triassic flora from Southwest China. In PDF, Science Bulletin, 63: 1462–1463.

D.K. Ferguson (1967): On the phytogeography of coniferales in the European cenozoic. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 3: 73-110. See also here.

! Florida Museum of Natural History, University of Florida, Gainesville:
Fossil coniferophytes (Powerpoint presentatation).

Debbie Folkerts, Auburn University, Auburn, Alabama: Kingdom Plantae: Gymnosperms. Powerpoint presentation.
The link is to a version archived by the Internet Archive´s Wayback Machine.

G. Forte et al. (2022): Conifer Cone and Dwarf Shoot Diversity in the Anisian (Middle Triassic) of Kühwiesenkopf/Monte Prà della Vacca (Dolomites, Northeastern Italy). Abstract, International Journal of Plant Sciences, 183.

G. Forte et al. (2021): Conifer Diversity in the Middle Triassic: New Data from the Fossillagerstätte Kühwiesenkopf/Monte Prà della Vacca (Pelsonian, Anisian) in the Dolomites (Northeastern Italy). Abstract, Int. J. Plant Sci., 182: 445–467.

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

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

Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine: Gymnosperms in the Mesophytic.
The link is to a version archived by the Internet Archive´s Wayback Machine.

D.S. Gernandt et al. (2018): Incorporating fossils into the Pinaceae tree of life. Open access, American Journal of Botany, 105: 1329–1344.

D.S. Gernandt et al. (2016): Phylogenetics of extant and fossil Pinaceae: methods for increasing topological stability. Abstract, Botany, 94: 863-884. See also here (in PDF).

D.S. Gernandt et al. (2008): Use of simultaneous analyses to guide fossil-based calibrations of Pinaceae phylogeny. PDF file, Int. J. Plant Sci., 169: 1086-1099.

S. Gilmore and K.D. Hill (1997): Relationships of the Wollemi Pine (Wollemia nobilis) and a molecular phylogeny of the Araucariaceae. PDF file, Telopea 7. See also here.

S.C. Gnaedinger and A.M. Zavattieri (2017): First Record of Voltzialean Male Cone (Lutanthus) and Podocarpacean Female Cone (Rissikistrobus) from the Late Triassic of Argentina, Including New Plant Remains from the Paso Flores Formation. Abstract, Ameghiniana, 54: 224–246. See also here (in PDF).

U.G. Hacke et al. (2015): The Hydraulic Architecture of Conifers in Ecological and Functional Xylem Anatomy. In PDF, book chapter, Springer International.

M. Hámor-Vidó et al. (2010): In situ preservation and paleoenvironmental assessment of Taxodiacea fossil trees in the Bükkalja Lignite Formation, Bükkábrány open cast mine, Hungary. In PDF, International Journal of Coal Geology, 81: 203–210.
See also here.
Note figure 2: Fossil trees are standing on the top of the top of lignite seam.

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.

T. He et al. (2016): A 350-million-year legacy of fire adaptation among conifers. Abstract, Journal of Ecology, 104: 352–363. See also here (in PDF).

T. He et al. (2012): Fire-adapted traits of Pinus arose in the fiery Cretaceous. Free access, New Phytologist, 194: 751–759. See also here (in PDF).

R. Heady (2012): The Wollemi Pine—16 years on. In PDF, Chapter 15: Australia’s Ever-changing Forests VI: Proceedings of the Eighth National Conference on Australian Forest History. Brett J. Stubbs et al. (ed.).
Snapshot provided by the Internet Archive´s Wayback Machine.

R.D. Heady and G.E. Burrows (2008): Features of the secondary xylem that facilitate branch abscission in juvenile Wollemia nobilis. In PDF, IAWA Journal, 29: 225-236.

E.J. Hermsen et al. (2007): A voltzialean pollen cone from the Triassic of Antarctica. In PDF, Review of Palaeobotany and Palynology, 144: 113-122.
The link is to a version archived by the Internet Archive´s Wayback Machine.

F. Herrera et al. (2020): Reconstructing Krassilovia mongolica supports recognition of a new and unusual group of Mesozoic conifers. Open access, PLoS ONE, 15: e0226779.
Note figs 6, 7: Reconstructions of Krassilovia mongolica. Drawings: Pollyanna von Knorring.

F. Herrera et al. (2015): A New Voltzian Seed Cone from the Early Cretaceous of Mongolia and Its Implications for the Evolution of Ancient Conifers. In PDF, Int. J. Plant Sci., 176: 791-809.

! J. Herting and T. Stützel (2022): Evolution of the coniferous seed scale. Free access, Annals of Botany, 129: 753–760.
Note figure 1: Expanded derivation series of extant conifer seed scale morphology.
Figure 2: Possible ways of modification of the funiculus in taxa with a morphology like Pinaceae and Cunninghamia.

T.J. Hieger et al. (2015): Cheirolepidiaceous diversity: An anatomically preserved pollen cone from the Lower Jurassic of southern Victoria Land, Antarctica. In PDF, Review of Palaeobotany and Palynology, 220: 78–87. See also here.

T.E. Higham et al. (2022): The Evolution of Mechanical Properties of Conifer and Angiosperm Woods. In PDF, Integrative and Comparative Biology, 62: 668–682.
See also here.

! J. Hilton et al. (2016): Age and identity of the oldest pine fossils: COMMENT. Geology, 44. See also:
! Reaffirming Pinus mundayi as the oldest known pine fossil: REPLY. By H.J. Falcon-Lang et al., 2016.
Please take notice:
The oldest Pinus and its preservation by fire. Abstract, by H.J. Falcon-Lang et al., 2016.

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

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

Armin Jagel (2001): Morphologische und morphogenetische Untersuchungen zur Systematik und Evolution der Cupressaceae s. l. (Zypressengewächse). PDF file (46 MB), in German. Dissertation, Ruhr-Universität Bochum.
A version archived by the Internet Archive´s Wayback Machine.
See also here.

Ross Koning, Biology Department, Eastern Connecticut State University, Willimantic: Biology of Plants. This course is an introduction to botany and stresses anatomy, morphology, natural history, and evolution among organisms called plants. Go to: The Naked Seeds of Pinus. See also here.
All links are available through the Internet Archive´s Wayback Machine.

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 et al. (2013): Aberrant Classopollis pollen reveals evidence for unreduced (2n) pollen in the conifer family Cheirolepidiaceae during the Triassic-Jurassic transition. Free access, Proc. R. Soc. B, 280.

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

A.B. Leslie and R.B.J. Benson (2022): Neontological and paleontological congruence in the evolution of Podocarpaceae (coniferales) reproductive morphology. Free access, Front. Ecol. Evol., 10: 1058746. doi: 10.3389/fevo.2022.1058746
"... Although molecular and fossil data regarding the deep evolution of Podocarpaceae reproductive structures are sparse, they are complementary. Both suggest that cones, seeds, and leaves evolved as a suite of traits in a stepwise manner in response to changing ecological conditions from the Early Cretaceous through the early Cenozoic ..."

A.B. Leslie and J.M. Losada (2019): Reproductive Ontogeny and the Evolution of Morphological Diversity in Conifers and Other Plants. Free access, Integrative and Comparative Biology, 59: 548–558.
Note figure 4: Depictions of ovuliferous complex and ovule/seed ontogeny in different conifer groups through time.

A.B. Leslie et al. (2018): An overview of extant conifer evolution from the perspective of the fossil record. Abstract, American Journal of Botany, 105: 1–14. See also here (in PDF).

! A.B. Leslie et al. (2015): Integration and macroevolutionary patterns in the pollination biology of conifers. In PDF, Evolution, 69: 1573-1583.

! A.B. Leslie et al. (2012): Hemisphere-scale differences in conifer evolutionary dynamics. In PDF, PNAS, 109: 16217-16221. See also here.

Gerhard Leubner Lab, University Freiburg, Germany: Seed Evolution. Go to: Conifers - Life cycle of pine: Extant gymnosperms, the oldest trees .

Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima.
This expired link is now available through the Internet Archive´s Wayback Machine.

A. Linkies et al. (2010): The evolution of seeds. PDF file, New Phytologist.

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

J.M. Losada et al. (2019): Not all 'pine cones' flex: functional trade-offs and the evolution of seed release mechanisms. Free access, New Phytologist, 222: 396–407.

Y. Lu et al. (2013): Determination of the molecular signature of fossil conifers by experimental palaeochemotaxonomy – Part 1: The Araucariaceae family. Biogeosciences, 10, 1943–1962,

Y. Lu et al. (2012): Determination of the molecular signature of fossil conifers by experimental palaeochemotaxonomy - Part 1: The Araucariaceae family. In PDF, Biogeosciences Discuss., 9: 10513-10550.

Y. Lu et al. (2011): Adaptation of male reproductive structures to wind pollination in gymnosperms: Cones and pollen grains. In PDF, Can. J. Plant Sci. 91: 897-906. See also here.

J. Ma (2003): The chronology of the "living fossil" Metasequoia glyptostroboides (Taxodiaceae): a review (1943-2003). PDF file, Harvard Papers in Botany, 8: 9-18. See also here.
"... On the basis of primary documents including letters, manuscripts, and original publications, plus personal experience, the major events, important publications, and main scientists related to this story are recorded chronologically for the first time 60 years after the species’ discovery ..."

Department of Botany, University of Wisconsin, Madison: Plant Systematics Collection. This web site provides structured access to a teaching collection of plant images representing over 250 families and 1000 genera of vascular plants. Go to: Phylum Coniferophyta (The Conifers).
These expired links are now available through the Internet Archive´s Wayback Machine.

K. Mao et al. (2012): Distribution of living Cupressaceae reflects the breakup of Pangea. In PDF, Proc. Natl. Acad. Sci., 109: 7793-7798.

J. Marmi et al. (2023): Evolutionary history, biogeography, and extinction of the Cretaceous cheirolepidiaceous conifer, Frenelopsis. Free access, Evolving Earth, 1.

L. Marynowski et al. (2007): Biomolecules preserved in ca. 168 million year old fossil conifer wood. PDF file, Naturwissenschaften, 94: 228-236.

K.K.S. Matsunaga et al. (2021): Ovulate Cones of Schizolepidopsis ediae sp. nov. Provide Insights into the Evolution of Pinaceae. Free access, Int. J. Plant Sci., 182: 490–507.

C. Mays et al. (2017): Polar wildfires and conifer serotiny during the Cretaceous global hothouse. In PDF, Geology, 45: 1119-1122. See also here.

C. Mays et al. (2017): Pushing the limits of neutron tomography in palaeontology: Three-dimensional modelling of in situ resin within fossil plants. Open access, Palaeontologia Electronica, 20.3.57A: 1-12.
See also here.
"... This study demonstrates the feasibility of NT [neutron tomography] as a means to differentiate chemically distinct organic compounds within fossils ..."

! S. McLoughlin (2021): Gymnosperms: History of Life: Plants: Gymnosperms. PDF file, in: Elias, S. and Alderton, D. (eds): Encyclopedia of Geology. See also here.

M.M. Mendes et al. (2018): Some Conifers from The Early Cretaceous (Late Aptian–Early Albian) of Catefica, Lusitanian Basin, Western Portugal. Free access, Fossil Imprint, 74: 317–326.

D. Mietchen et al. (2008): Three-dimensional Magnetic Resonance Imaging of fossils across taxa PDF file, Biogeosciences, 5: 25-41. Fossil cones of the conifer Pararaucaria patagonica. See also here.

M. Muir and J.H.A. van Konijnenburg-van Cittert (1970): A Rhaeto-Liassic flora from Airel, Northern France. In PDF.

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, CONIFERS. Link list page with rankings and brief explanations. Images of Lebachia, Walchia, Walchia piniformis, Cassinisia orobica, Pseudovoltzia liebeana, Majonica alpina, Dolomitia cittertiae.
Snapshot provided by the Internet Archive´s Wayback Machine.

N. Nestle et al. (2018): Fossilized but functional – Tomographic insights into nature’s most resilient actuators. In PDF, Micro-CT User Meeting.
See also here (P. Gaupels, Geohorizon; in German).

Dan Nickrent and Karen Renzaglia, Department of Plant Biology, Southern Illinois University at Carbondale: Land Plants Online, Phylum Pinophyta, The Conifers (now via wayback archive).

S. Nigris et al. (2021): Fleshy Structures Associated with Ovule Protection and Seed Dispersal in Gymnosperms: A Systematic and Evolutionary Overview. Open access, Critical Reviews in Plant Sciences, 40.

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

C.A. Offord et al. (1999): Sexual Reproduction and Early Plant Growth of the Wollemi Pine (Wollemia nobilis), a Rare and Threatened Australian Conifer. PDF file, Annals of Botany 84.

A. Otto and B.R.T. Simoneit (2001): Chemosystematics and diagenesis of terpenoids in fossil conifer species and sediment from the Eocene Zeitz formation, Saxony, Germany. In PDF, Geochimica et Cosmochimica Acta, 65: 3505-3527.
See also here.

Osborne, C.P. & Beerling, D.J. (2002): A process-based model of conifer structure and function with special emphasis on leaf lifespan.. PDF file, Global Biogeochemical Cycles.
Now provided by the Internet Archive´s Wayback Machine.

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

G. Pacyna et al. (2017): A new conifer from the Upper Triassic of Southern Poland linking the advanced voltzialean type of ovuliferous scale with Brachyphyllum/Pagiophyllum-like leaves. Abstract, Review of Palaeobotany and Palynology, 245: 28-54. See also here (in PDF).

G.A. Pattemore and A.C. Rozefelds (2019): Palissya – absolutely incomprehensible or surprisingly interpretable: a new morphological model, affiliations and phylogenetic insights. Open access, Acta Palaeobotanica, 59: 181–214.

G.A. Pattemore et al. (2015): Palissya: A global review and reassessment of Eastern Gondwanan material. In PDF, Review of Palaeobotany and Palynology, 210: 50-61.

V. Pietzsch and V.M. Dörken (2018): Morphological and anatomical investigation of seed cones of Cupressus glabra (Cupressaceae): evolutionary aspects. In PDF, Bull. Cupressus Conservation Proj., 7: 82-91.

Kathleen B. Pigg, Department of Plant Biology, Arizona State University:
Plant Fossils and Evolution.
! Go to: Laboratory 11. Paleozoic Seed Ferns, Cordaites & Early Conifers, Gondwana groups.
Websites outdated. Links lead to versions archived by the Internet Archive´s Wayback Machine.

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

M. Pole et al. (2016): The rise and demise of Podozamites in east Asia - An extinct conifer life style. Abstract. See also here.

S. Poppinga et al. (2016): Hygroscopic motions of fossil conifer cones. Scientific Reports, 7.

Ruud J. Poort, Henk Visscher, and David L. Dilcher: Zoidogamy in fossil gymnosperms: The centenary of a concept, with special reference to prepollen of late Paleozoic conifers. The National Academy of Sciences, PNAS 1996 93: 11713-11717.

! 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.

! S. Renner (2009): Gymnosperms. Provided by the Internet Archive´s Wayback Machine. PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

J. Restemeyer, Fakultät für Biologie, Ruhr-Universität Bochum: Morphologische und morphogenetische Untersuchungen zur Phylogenie und Evolution der Podocarpaceae und Phyllocladaceae. Thesis, in German.

David M. Richardson (ed.; 1998): Ecology and Biogeography of Pinus. 546 pages, Google books.
See also here.
Book review.

! G. Roghi et al. (2017): Middle Triassic amber associated with voltzialean conifers from the Southern Alps of Italy. Abstract, Rivista Italiana di Paleontologia e Stratigrafia, 123: 193-202. See also here (in PDF).

R. Rößler et al. (2014): Which name(s) should be used for Araucaria-like fossil wood?—Results of a poll. In PDF, Taxon, 63: 177–184. See also here.

G.W. Rothwell and T. Ohana (2016): Stockeystrobus gen. nov. (Cupressaceae), and the evolutionary diversification of sequoioid conifer seed cones. Abstract, Botany, 94: 847-861. See also here (in PDF).

G.W. Rothwell et al. (2013): Diversity of Ancient Conifers: The Jurassic Seed Cone Bancroftiastrobus digitata gen. et sp. nov.(Coniferales). In PDF, International Journal of Plant Sciences, 174: 937-946.

G.W. Rothwell et al. (2012): The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae. In PDF, American Journal of Botany, 99: 708–720.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens, OH: Vascular Plant Morphology. Archived by Internet Archive Wayback Machine. This course covers the structure, development, reproductive biology and relationships of vascular plants. The course is structured to emphasize the evolutionary changes that led to the diversity of modern tracheophytes. Go to Cordaitales and Coniferales (PDF file).

G.W. Rothwell et al. (2005): Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). PDF file, Taxon 54: 733–750.
See also here.

D.P. Ruiz and J. Bodnar (2019): The oldest record of Juniperoxylon, a cupressaceous fossil wood from the Middle Triassic of Argentina. In PDF, Acta Palaeontologica Polonica, 64: 481–488.

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

B. Saladin et al. (2017): Fossils matter: improved estimates of divergence times in Pinus reveal older diversification. Open Access, BMC Evolutionary Biology.

von Schauroth (1852): Ueber das Vorkommen von Voltzia coburgensis im mittleren Keupersandstein- Zeitschrift der Deutschen geologischen Gesellschaft, 4: 538-544. Provided by

C. Schulz et al. (2014): Male Cone Evolution in Conifers: Not All That Simple. In PDF, American Journal of Plant Sciences, 5: 2842-2857. See also here and there.

E. Schütze (1901): Beiträge zur Kenntnis der triassischen Koniferengattungen: Pagiophyllum, Voltzia und Widdringtonites. PDF file, in German; Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg, 57: 240-273, plate VI bis X, Stuttgart.
See also here.

A.B. Schwendemann et al. (2011): Morphological and functional stasis in mycorrhizal root nodules as exhibited by a Triassic conifer. In PDF.

! A.B. Schwendemann et al. (2010): Organization, anatomy, and fungal endophytes of a Triassic conifer embryo. Open access, American Journal of Botany, 97: 1873-1883.

! R. Serbet et al. (2013): Cunninghamia taylorii sp. nov., a Structurally Preserved Cupressaceous Conifer from the Upper Cretaceous (Campanian) Horseshoe Canyon Formation of Western North America. In PDF, International Journal of Plant Sciences, 174: 471-488. See also here.

A.S. Sequeira and B.D. Farrell (2001): Evolutionary origins of Gondwanan interactions: How old are Araucaria beetle herbivores? PDF file, Biological Journal of the Linnean Society 74: 459-474. See also here.
This expired link is still available through the Internet Archive´s Wayback Machine.

G. Shi et al. (2014): Whole-Plant Reconstruction and Phylogenetic Relationships of Elatides zhoui sp. nov. (Cupressaceae) from the Early Cretaceous of Mongolia. In PDF, International Journal of Plant Sciences, 175. See also here.

K.C. Shunn and C.T. Gee (2023): Cross-sectioning to the core of conifers: pith anatomy of living Araucariaceae and Podocarpaceae, with comparisons to fossil pith. Open access, IAWA Journal.
"... In addition to a general paucity in pith descriptions [...] we focus here on the pith of 16 conifer species [...] as well as comparing pith anatomy in regard to branch age, genus, and family. Furthermore, comparisons are made to fossil conifer pith to elucidate common features shared by living conifers and their ancient relatives ..."

Thomas Speck and Sylke Döringhoff, Botanical Garden, Freiburg University, Germany: Coniferetum (in German).
This expired link is available through the Internet Archive´s Wayback Machine.

! A.R.T. Spencer et al. (2015): Middle Jurassic evidence for the origin of Cupressaceae: A paleobotanical context for the roles of regulatory genetics and development in the evolution of conifer seed cones. Free access, American Journal of Botany, 102: 942-961.

D.C. Steart et al. (2014): X-ray Synchrotron Microtomography of a silicified Jurassic Cheirolepidiaceae (Conifer) cone: histology and morphology of Pararaucaria collinsonae sp. nov. In PDF, see also here.

R.A. Stockey et al. (2020): Late Cretaceous Diversification of Cupressaceous Conifers: A Taiwanioid Seed Cone from the Eden Main, Vancouver Island, British Columbia, Canada. In PDF, International Journal of Plant Sciences 181. See also here.

! R.A. Stockey and G.W. Rothwell (2020): Diversification of crown group Araucaria: the role of Araucaria famii sp. nov. in the Late Cretaceous (Campanian) radiation of Araucariaceae in the Northern Hemisphere. Abstract, American Journal of Botany, 107: 1–22. See also here (in PDF).

R.A. Stockey (1981): Some comments on the origin and evolution of conifers Canadian Journal of Botany, 59: 1932-1940. See also here.

R.A. Stockey (1977): Reproductive biology of the Cerro Cuadrado (Jurassic) fossil conifers: Pararaucaria patagonica. In PDF, American Journal of Botany, 64: 733-744. See also here.

Ed Strauss, Washington (article hosted by Evolving Earth Foundation Issaquah, WA). The Evolving Earth Foundation is committed to encouraging research and building community related to the earth sciences.
How to Identify Conifers. Conifer micro photographs.
Websites still available via Internet Archive Wayback Machine.

M. Sundaram et al. (2019): Accumulation over evolutionary time as a major cause of biodiversity hotspots in conifers. In PDF, Proc. R. Soc. B, 286: 20191887. See also here.

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
! BOT335 Lecture Schedule. Some interesting chapters in terms of palaeobotany, e.g.
The First Vascular Land Plants;
Carboniferous Forests;
Arborescent Lycopods;
Psaronius: a Carboniferous tree-fern;
Carboniferous Horsetails;
Carboniferous Seed Ferns;
The Evolution of Conifers;
Cycadophytes, the True Cycads;
Mesozoic Cycadeoids;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

M.L. Trivett and G.W. Rothwell (1991): Diversity among Paleozoic Cordaitales. In PDF, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 183: 289-305.

M.L. Trivett and G.W. Rothwell (1985): Morphology, systematics, and paleoecology of Paleozoic fossil plants: Mesoxylon priapi, sp. nov.(Cordaitales). In PDF, Systematic Botany, 10: 205-223.
See also here.

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

K.Y. Wang et al. (2022): Anatomically preserved cordaitalean trees from the Pennsylvanian of Yangquan City, Shanxi Province, and their implication for a perhumid climate in North China Block. In PDF, Palaeoworld, 31: 294-310.
See also here.

Biology Department, Western Washington University, Bellingham, Washington:
! Naked seeded vascular plants (Conifers, etc.) Powerpoint presentation. See also here, or there.

! Wikipedia (a free-content encyclopedia): Spermatophyte. Go to: The conifers.

Wikipedia, the free encyclopedia: Wollemia.

J.P. Wilson and A.H. Knoll (2010): A physiologically explicit morphospace for tracheid-based water transport in modern and extinct seed plants. PDF file, Paleobiology, 36: 335-355.
See also here.

C. Witkowski (2014): Mimicking Early Stages Of Diagenesis In Modern Metasequoia Leaves Implications For Plant Fossil Lagerstätten. In PDF, Thesis in Global Environmental Studies, Department of Science and Technology, Bryant University (Master of Science in Global Environmental Studies).
See also here. Abstract, Session No. 17: An Interdisciplinary Approach to Taphonomy: The Impact of Morphological, Molecular, and Isotopic Changes on Environmental Proxies. Northeastern Section, 49th Annual Meeting, The Geological Society of America.

Wollemi Australia Pty Ltd (a subsidiary company of Forestry Plantations Queensland, the principal commercial plantation forest grower in Queensland, Australia): The Wollemi Pine. One of the world's oldest and rarest plants dating back to the Cretaceous. See also: Gallery (in German).

Wollemi Pine North America, San Diego, CA: About Wollemi.
This expired link is available through the Internet Archive´s Wayback Machine.

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

X. Yao et al. (1993): The triassic seed cone Telemachus from Antarctica. PDF file, Review of palaeobotany and palynology, 78: 269-276.
See also here.

Jian-Wei Zhang et al. (2012): A new species of the extinct genus Austrohamia (Cupressaceae s.l.) in the Daohugou Jurassic flora of China and its phytogeographical implications. In PDF, Journal of Systematics and Evolution, 50: 72-82. See also here (abstract).

L. Zhang et al. (2021): First fossil foliage record in the red beds from the Upper Jurassic in the Sichuan Basin, southern China. In PDF, Geological Journal. See also here.
Note fig. 6: Comparisons with potential Mesozoic conifer fossils.

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Last updated September 16, 2023

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