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

! J. Asar et al. (2022): Early diversifications of angiosperms and their insect pollinators: were they unlinked? Free access. Trends in Plant Science, 27: 858-869. See also here.
Note figure 1: Emergence of crown angiosperms and insect pollinators.
Figure 2. Phylogeny of seed plants, depicting pollination modes of both extinct and extant lineages.

! R.M. Bateman et al. (2006): Morphological and molecular phylogenetic context of the angiosperms: contrasting the ‘top-down’ and ‘bottom-up’ approaches used to infer the likely characteristics of the first flowers. Free access, Journal of Experimental Botany, Vol. 57, No. 13, pp. 3471–3503. Major Themes in Flowering Research Special Issue.
Note fig. 1C: Male and female reproductive structures of Caytonia (Caytoniales).
Fig. 1D: Reproductive structure of Williamsoniella (Bennettitales).

Museum of Paleontology (UCMP), University of California at Berkeley: Introduction to the Bennettitales, the cycadeoids.

R. Barboni and T.L. Dutra (2013): New "flower" and leaves of Bennettitales from Southern Brazil and their implication in the age of the Lower Mesozoic deposits. In PDF, Ameghiniana, 50: 14-32.

P. Blomenkemper et al. (2021): Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group. In PDF, Front. Earth Sci., 9: 652699. doi: 10.3389/feart.2021.652699.
See also here.

Philippe Choler, Laboratoire de Biologie des Populations d'Altitude, Université Joseph Fourier, Grenoble: Biologie Evolutive Végétale. Snapshot taken by the Internet Archive´s Wayback Machine. Concepts and methods in evolutionary biology (in French). Navigate from "Plan du cours" (access to about 335 slides). Go to: Bennettitales (1), and Bennettitales (2).

C.J. Cleal et al. (2006): A clarification of the type of the morphogenus Nilssoniopteris Nathorst (Gymnospermophyta, Bennettitales, fossiles). PDF file, Taxon, 55: 219-222.

! P.R. Crane and P.S. Herendeen (2009): Bennettitales from the Grisethorpe Bed (Middle Jurassic) at Cayton Bay, Yorkshire, UK. Open access, American Journal of Botany, 96: 284-295.

! N.R. Cúneo et al. (2010): Review of the Cycads and Bennettitaleans from the Mesozoic of Argentina. In PDF.

Z. Czier (2014): A fern-bennettitalean floral assemblage in Tithonian-Berriasian travertine deposits (Aguilar Formation, Burgos-Palencia, N Spain) and its palaeoclimatic and vegetational implications. In PDF, Journal of Iberian Geology, 35: 127-140.
Specimens preserved as impressions coated with a microbial film up to 5 mm thick made up of bacteria and cyanobacteria.

! A.B. Doweld (2016): Proposals to conserve the names Cycadeoidea against Mantellia and C. megalophylla against M. nidiformis (fossil Spermatophyta: Cycadeoideopsida). In PDF, Taxon, 65: 397-399.

H. El Atfy et al. (2022): Zamites (Bennettitales) from the Minjur Formation (Norian) of Saudi Arabia – a unique record from the Late Triassic palaeotropics of Gondwana. Full access, Botany Letters, DOI: 10.1080/23818107.2022.2088614.
See also here (in PDF).

D. Frame and G. Gottsberger (2023): Diverse sexual strategies in fossil gymnosperms: pollination in the Bennettitales revisited. In PDF, Phyton, 62–63: 127–137. See also here.
"... Our review grounded in modern concepts of floral biology and plant-animal interactions leads to new interpretations of existing data
[...] Pollination was similar to an angiosperm cantharophilous syndrome, complete with pollination chamber, except that bisexual bennetittalean flowers were protandrous rather than protogynous
[...] Beetles in the pollination chamber mated and females oviposited in the androecium ..."

! E.M. Friis et al. (2013): New Diversity among Chlamydospermous Seeds from the Early Cretaceous of Portugal and North America. Free accesss, International Journal of Plant Sciences, 174: 530–558.
"... The material is based on numerous charcoalified and lignitic specimens recovered from Early Cretaceous mesofossil floras [...]
! Attenuation-based synchrotron-radiation x-ray tomographic microscopy (SRXTM) and phase-contrast x-ray tomographic microscopy (PCXTM) were carried out [...]
! Volume rendering (voltex), which provides transparent reconstructions, was also used for the virtual sections ..."

! E.M. Friis et al. (2011): Early Flowers and Angiosperm Evolution. Abstract, Cambridge University Press.
See also here (in PDF, long download time) and there (Google books). See especially chapter 5: Angiosperms in context: extant and fossil seed plants
Also worth to check out: Book Review, by P.J. Rudall, Botanical Journal of the Linnean Society, 170. In PDF.
"... This long-awaited book represents not only a remarkable tour de force of palaeobotanical literature, but also a potentially enduring biological textbook. ..."

Else Marie Friis et al. (2007): Phase-contrast X-ray microtomography links Cretaceous seeds with Gnetales and Bennettitales. Abstract, Nature 450: 549-552.
! See also here (in PDF).

! M.W. Frohlich & M.W. Chase (2007): After a dozen years of progress the origin of angiosperms is still a great mystery. In PDF, Nature, 450: 1184-1189.
See also here.

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

D.S. Guzmán and M.P. Velasco (2021): Weltrichia magna sp. nov., a new record for the Middle Jurassic of Oaxaca, Mexico. In PDF, Acta Palaeobotanica, 61: 95–106.
Note fig. 2: Reconstructions proposed for Weltrichia magna.

T.M. Harris (1973): The strange Bennettitales. In PDF, Sir Albert Charles Seward Memorial Lecture, Birbal Sahni Institute of Palaeobotany.
This expired link is available through the Internet Archive´s Wayback Machine.

R. Herbst and A. Crisafulli (2016): Buckya austroamericana nov. gen. et sp. (Bennettitales) from the Upper Triassic Laguna Colorada Formation (El Tranquilo Group), Santa Cruz province, Argentina. In PDF, Serie Correlación Geológica, 32: 85-100. See also here.

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

Valentin A. Krassilov (1987): Palaeobotany of the mesophyticum: state of the art. In PDF, Review of Palaeobotany and Palynology, 50: 231-254. Provided by the Internet Archive´s Wayback Machine.

! C.C. Labandeira et al. (2016): The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies. See also here (in PDF). Proc. R. Soc., B 283.
Heritagedaily: Paleobotanist plays role in discovery of "Jurassic butterflies". An artist´s rendering of the butterfly Oregramma illecebrosa, consuming pollen drops from Triassic bennettitales.

Gerhard Leubner Lab, University Freiburg, Germany: Seed Evolution. Go to: Jurassic parc/Mesozoic era: Extinct gymnosperms and living ancient gymnosperms. Extant Cycadales and extinct Bennettitales.

F. Liu et al. (2022): Anatomical Study of Cretaceous, Permineralized, Bennettitalean Fossils from Heilongjiang Province, NE China. Free access, Acta Geologica Sinica.

Z.J. Liu et al. (2019): Zhangwuia: an enigmatic organ with a bennettitalean appearance and enclosed ovules. In PDF, Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 108 (Agora Paleobotanica), : 419-428. See also here.

D.E. Lozano-Carmona et al. (2019): Jurassic Flora in Southern Mexico: Anomozamites Schimper, 1870 Emend. Pott et McLoughlin, 2009 from Mixteco Terrane, Phytogeographical Implications of Williamsoniaceae Family (Bennettitales). Open Journal of Geology, 9: 142-156.

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

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

John M. Miller (gigantopteroid.org), University of California, Berkeley: Origin of Angiosperms. See also here or navigate from essay contents.
These expired links are now available through the Internet Archive´s Wayback Machine.

! D. Naware et al. (2024): Patterns of variation in fleshy diaspore size and abundance from Late Triassic–Oligocene. Open access, Biological Reviews, 99: 430-457.
Note figure 2: Genus richness of seed plants with fleshy and non-fleshy diaspores from Late Triassic to Oligocene across mid- to high latitudes and low latitudes.
"... Vertebrate-mediated seed dispersal is a common attribute of many living plants, and variation in the size and abundance of fleshy diaspores is influenced by regional climate and by the nature of vertebrate seed dispersers among present-day floras.
[...] We present a new data set of more than 800 georeferenced fossil diaspore occurrences spanning the Triassic–Oligocene, across low to mid- to high palaeolatitudes ..."

The Palm and Cycad Society of Florida:
The fossil Cycads. With paintings and reconstructions of Douglas Henderson, John Sibbick, and Mark Hallett.
Go to: Cretaceous Cycadales. Reconstruction images, image of Cycadeoidea marylandica.
Websites still available through the Internet Archive´s Wayback Machine.

! E. Peñalvera et al. (2012): Thrips pollination of Mesozoic gymnosperms. In PDF, PNAS, 109: 8623-8628. See also here.

! Kathleen B. Pigg, Department of Plant Biology, Arizona State University: Laboratory. Go to: The Cycads, Cycadeoids (Bennettitales) and Ginkgophytes.

C. Pott and J.H.A. Van Konijnenburg-Van Cittert (2017): The type specimen of Nilssoniopteris solitaria (Phillips 1829) (Bennettitales). Open access, Acta Palaeobotanica, 57: 177–184.

C. Pott et al. (2017): Lunzia austriaca – a bennettitalean microsporangiate structure with Cycadopites-like in situ pollen from the Carnian (Upper Triassic) of Lunz, Austria. Abstract, Grana, 56. See also here (in PDF).
Depicted in fig. 8: Restoration of the cup-shaped Lunzia microsporangiate organ as interpreted from the fossils.

C. Pott et al. (2016): Bennettitales in the Rhaetian flora of Wüstenwelsberg, Bavaria, Germany. Abstract, Review of Palaeobotany and Palynology, 232: 98–118. See also here (in PDF).

C. Pott (2016): Westersheimia pramelreuthensis from the Carnian (Upper Triassic) of Lunz, Austria: More Evidence for a Unitegmic Seed Coat in Early Bennettitales. In PDF, International Journal of Plant Sciences.
See also here.

C. Pott and B.J. Axsmith (2015): Williamsonia carolinensis sp. nov. and Associated Eoginkgoites Foliage from the Upper Triassic Pekin Formation, North Carolina: Implications for Early Evolution in the Williamsoniaceae (Bennettitales). In PDF, International Journal of Plant Sciences, Vol. 176: 174-185.

C. Pott et al. (2015): Wielandiella villosa comb. nov. from the Middle Jurassic of Daohugou, China: More Evidence for Divaricate Plant Architecture in Williamsoniceae. In PDF, Botanica Pacifica, 4.

C. Pott and S. McLoughlin (2014): Divaricate growth habit in Williamsoniaceae (Bennettitales): unravelling the ecology of a key Mesozoic plant group. Abstract, Palaeobiodiversity and Palaeoenvironments, 94: 307–325. See also here (in PDF).

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

! C. Pott et al. (2012): Baikalophyllum lobatum and Rehezamites anisolobus: Two Seed Plants with "Cycadophyte" Foliage from the Early Cretaceous of Eastern Asia. In PDF, International Journal of Plant Sciences, 173: 192-208.
See likewise here.
Paper awarded with the Remy and Remy Award 2012, Botanical Society of America.

! 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.

Christian Pott, Natural History Museum Stockholm: Bennettitaleans of the world. The aim of this website is to compile as much information as possible about this enigmatic group of gymnosperms. Under construction.

! C. Pott et al. (2010): Late Palaeozoic foliage from China displays affinities to Cycadales rather than to Bennettitales necessitating a re-evaluation of the Palaeozoic Pterophyllum species. PDF file, Acta Palaeontol. Pol., 55: 57-168.

C. Pott and M. Krings (2007): First record of circinate vernation in bennettitalean foliage. In PDF, N. Jb. Geol. Paläont. Abh., 245: 315-321. see also here (abstract).

C. Pott et al. (2007): Proposal to conserve the name Pterophyllum (Foss., Bennettitales) with a conserved type. PDF file, Taxon, 56: 966-967.

! S.E. Pratt (2014): Benchmarks: September 1, 1957: Fossil Cycad National Monument is dissolved. Earth, September 01, 2014.
See also here (National Fossil Day).
These expired links are now available through the Internet Archive´s Wayback Machine.

Dong Ren et al. (2009): A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies. PDF file, Science, 326: 840-847. See also here.

! G.W. Rothwell et al. (2009): Is the anthophyte hypothesis alive and well? New evidence from the reproductive structures of Bennettitales. Free access, American Journal of Botany, 96: 296-322. Note fig. 1: Cycadeoidea spp. Characteristic features of Cycadeoidea plants.
! Table 2. Contrasting characters of Bennettitales and Cycadales.

G.W. Rothwell and R.A. Stockey (2002): Anatomically preserved Cycadeoidea (Cycadeoidaceae), with a reevaluation of systematic characters for the seed cones of Bennettitales. Free access, American Journal of Botany, 89: 1447–1458.

ROTHWELL, GAR W.¹ and RUTH STOCKEY². ¹Department of Environmental and Plant Biology, Ohio University, Athens; ²Department of Biological Sciences, University of Alberta, Edmonton: Systematic characters of ovulate Cycadeoidea/Bennettites cones from the Cretaceous of Vancouver and Hornby Islands, British Columbia, Canada. Abstract. Botany 2001, August 12 - 16, 2001; Albuquerque, New Mexico.

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

Vincent L. Santucci, National Park Service, Kemmerer, WY, and Marikka Hughes, Peabody Museum of Natural History, Paleobotany Division, New Haven: Fossil Cycad National Monument: A Case of Paleontological Resource Mismanagement. This expired link is available through the Internet Archive´s Wayback Machine.

Senckenberg Natural History Museum and Centre for Biodiversity Research, Frankfurt am Main: Entwicklung der Pflanzenwelt. Easy to understand introduction (in German). Image and reconstruction of Cycadeoidea. Now provided by the Internet Archive´s Wayback Machine.

Scott Singleton, Houston Gem and Mineral Society (2006): Cycad Anatomy and Fossil Occurrences in Texas. PDF file.

School of Ocean and Earth Science, Southampton Oceanography Centre, UK: Geology Collection, Plants. Provided by the Internet Archive´s Wayback Machine. Go to: Jurassic. Images of Cycadeoidea, Cycadeoidea gigantea, Cycadeoidea microphylla.

D. Soltis et al. (2017): Phylogeny and Evolution of the Angiosperms. Book announcement. See also here (Google books). Worth checking out:
! Relationships of Angiosperms to Other Seed Plants. In PDF.
Note figure 1.12: Reconstructions of Caytoniales.
Note figure 1.13: Reconstruction of Bennettitales.
Note figure 1.14: Reconstructions of Pentoxylon plants.
Note figure 1.15: Reconstructions of glossopterids.

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

Hans Steur, Ellecom, The Netherlands: The Jurassic flora of North Yorkshire, Bennettitales from Yorkshire.

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;
Ginkgophytes;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

L.B. Thien et al. (2000): New Perspectives on the Pollination Biology of Basal Angiosperms. Abstract, International Journal of Plant Sciences, 161.

J.H.A. van Konijnenburg-van Cittert et al. (2017): Differentiation of the fossil leaves assigned to Taeniopteris, Nilssoniopteris and Nilssonia with a comparison to similar genera, Abstract, Review of Palaeobotany and Palynology, 237: 100–106.See also here (in PDF).

! J.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 (after Cleal et al. 2001), how to distinguish bennettialean leaf shapes!
Now provided by the Internet Archive´s Wayback Machine.

M.P. Velasco-de León et al. (2024): New records of Bennettitales and associated flora from the Jurassic of the Cualac Formation, Mexico. Open access, Palaeontologia Electronica.

X. Wang et al. (2022): Nilssoniopteris longifolius Chang from the Middle–Late Jurassic of China: Implications for Bennettitales-insect interactions. In PDF, Review of Palaeobotany and Palynology, 297.
See also here.

Wikipedia, the free encyclopedia: Bennettitales.

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