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The Lost Clades – Extinct Bennettitales and Nilssoniales as a Key to Understanding the Origin and Evolution of Gymnosperms and Angiosperms


This project aims to understand the evolutionary history of the diverse, but poorly understood, complex of Mesozoic plants traditionally called “cycadophytes” and to clarify their potential as palaeoenvironmental and palaeoclimatic markers for the Mesozoic.

The study will be based on extensive museum collections of well-preserved leaves and reproductive organs supplimented by new field collections.

It will involve reconstruction of whole-plant-taxa that should serve as natural units for improved phylogenetic analyses, evaluation of the temporal and spatial diversity and diversification of cycadophytes to reveal evolutionary patterns in fertile and sterile structures (seed cones, ‘flowers’, foliage), and assessment of cuticular and anatomical adaptations that may be useful for interpreting local environmental to regional climatic conditions for these plants during the Mesozoic.

The major focus of the project will be on two enigmatic groups of extinct cycadophytes, i.e. the Bennettitales and the Nilssoniales.

This research is funded by the Swedish Research Council (VR)

External Project Participants

Brian Axsmith (researcher), University of South Alabama, Mobile, AL, USA

Jiang Baoyu (researcher), Nanjing University, Nanjing, PR China

Benjamin Bomfleur (researcher), University Münster, Germany

Guido Grimm (researcher), University Vienna, Austria

Hans Kerp (researcher), University Münster, Germany

Michael Krings (researcher), Bavarian State Collections, Munich, Germany

Jens Lehmann (researcher), University Bremen, Germany

Hideo Takimoto (researcher), Nature Museum Ibaraki, Japan

Johanna H. A. van Konijnenburg-van Cittert (professor emerita), Naturalis Leiden and University of Utrecht, The Netherlands

Selected Publications

Pott C, Fischer T, Aschauer B. Submitted. Lunzia austriaca — a bennettitalean reproductive structure with Cycadopites-like in situ pollen from the Carnian (Upper Triassic) of Lunz, Austria. Grana. DOI & PDF: not yet available

Van Konijnenburg-van Cittert JHA, Pott C, Cleal C, Zijlstra G. 2017. Differentiation of the fossil leaves assigned to Taeniopteris, Nilssoniopteris and Nilssonia with a comparison to similar genera. Review of Palaeobotany and Palynology. DOI & PDF: 10.1016/j.revpalbo.2016.11.009

Pott C. 2016. Westersheimia pramelreuthensis from the Carnian (Upper Triassic) of Lunz, Austria: More evidence for a unitegmic seed coat in early Bennettitales. International Journal of Plant Sciences 177: 771-791. DOI & PDF: 10.1086/688708

Pott C, Schmeissner S, Dütsch G, Van Konijnenburg-van Cittert JHA. 2016. Bennettitales in the Rhaetian flora of Wüstenwelsberg, Bavaria, Germany. Review of Palaeobotany and Palynology 232: 98-118. DOI & PDF: 10.1016/j.revpalbo.2016.04.010

Pott C, Axsmith BJ. 2015. Williamsonia carolinensis sp. nov. and associated Eoginkgoites foliage from the Upper Triassic Pekin Formation: Implications for early evolution in Williamsoniaceae (Bennettitales). International Journal of Plant Sciences 176: 174-185. DOI & PDF: 10.1086/679471

Pott C, Launis A. 2015. Taeniopteris novomundensis sp. nov. – “cycadophyte” foliage from the Carnian of Switzerland and Svalbard reconsidered: How to use Taeniopteris?. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 275: 19-31. DOI & PDF: 10.1127/njgpa/2015/0446

Pott C, Wang X, Zheng X. 2015. Wielandiella villosa comb. nov. from the Middle Jurassic of Daohugou, China: More evidence for divaricate plant architecture in Williamsoniaceae. Botanica Pacifica 4: 137-148. DOI & PDF: 10.17581/bp.2015.04215

Pott C. 2014. A revision of Wielandiella angustifolia – a shrub-sized bennettite from the Rhaetian–Hettangian of Scania, Sweden, and Jameson Land, Greenland. International Journal of Plant Sciences 175: 467-499. DOI & PDF: 10.1086/675577

Pott C, Guhl M, Lehmann J. 2014. The Early Cretaceous flora from the Wealden facies at Duingen, Germany. Review of Palaeobotany and Palynology 201: 75-105. DOI & PDF: 10.1016/j.revpalbo.2013.10.002

Pott C, McLoughlin S. 2014. Divaricate growth habit in Williamsoniaceae: Unravelling the ecology of a key Mesozoic plant group. Palaeobiodiversity and Palaeoenvironments 94: 307-325. DOI & PDF: 10.1007/s12549-014-0157-9

Pott C, McLoughlin S, Lindström A, Wu Shunqing, Friis EM. 2012. Baikalophyllum lobatum and Rehezamites anisolobus: Two seed plants with “cycadophyte” foliage from the Early Cretaceous of eastern Asia. International Journal of Plant Sciences 173: 192-208. DOI & PDF: 10.1086/663276

Pott C, McLoughlin S, Wu Shunqing, Friis EM. 2012. Trichomes on the leaves of Anomozamites villosus sp. nov. (Bennettitales) from the Daohugou beds (Middle Jurassic), Inner Mongolia, China: Mechanical defense against herbivorous arthropods? Review of Palaeobotany and Palynology 169: 48-60. DOI & PDF: 10.1016/j.revpalbo.2011.10.005

Bomfleur B, Pott C, Kerp H. 2011. Plant fossil assemblages of the Shafer Peak Formation (Lower Jurassic), north Victoria Land, East Antarctica. Antarctic Science 23: 188-208. DOI & PDF: 10.1017/S0954102011000149

McLoughlin S, Carpenter RJ, Pott C. 2011. Ptilophyllum muelleri (Ettingsh.) comb. nov. from the Oligocene of Australia – last of the Bennettitales? International Journal of Plant Sciences 172: 574-585. DOI & PDF: 10.1086/658920

Moisan P, Voigt S, Pott C, Buchwitz M, Schneider JW, Kerp H. 2011. Cycadalean and bennettitalean foliage from the Triassic Madygen Lagerstätte (SW Kyrgyzstan, Central Asia). Review of Palaeobotany and Palynology 164: 93-108. DOI & PDF: 10.1016/j.revpalbo.2010.11.008

Pott C, McLoughlin S. 2011. The Rhaetian flora from Rögla, northern Scania, Sweden. Palaeontology 54: 1025-1051. DOI & PDF: 10.1111/j.1475-4983.2011.01090.x

McLoughlin S, Pott C, Elliott D. 2010. The Winton Formation flora (Albian–Cenomanian, Eromanga Basin): Implications for vascular plant diversification and decline in the Australian Cretaceous. Alcheringa 34: 303-323. DOI & PDF: 10.1080/03115511003669944

Pott C, Krings M. 2010. Gymnosperm foliage from the Upper Triassic of Lunz, Lower Austria: An annotated checklist and identification key. Geo.Alp 7: 19-38. DOI not available. PDF for free

Pott C, Krings M, Kerp H, Friis EM. 2010. Reconstruction of a bennettitalean flower from the Carnian (Upper Triassic) of Lunz, Lower Austria. Review of Palaeobotany and Palynology 159: 94-111. DOI & PDF: 10.1016/j.revpalbo.2009.11.004

Pott C, McLoughlin S, Lindström A. 2010. Late Palaeozoic foliage from China displays affinities to Cycadales rather than to Bennettitales necessitating a re-evaluation of the Palaeozoic Pterophyllum species. Acta Palaeontologica Polonica 55: 157-168. DOI & PDF: 10.4202/app.2009.0070

McLoughlin S, Pott C. 2009. The Jurassic flora of Western Australia. GFF 131: 113-136. DOI & PDF: 10.1080/11035890902857846

Pott C, McLoughlin S. 2009. Bennettitalean foliage in the Rhaetian–Bajocian (latest Triassic–Middle Jurassic) floras of Scania, southern Sweden. Review of Palaeobotany and Palynology 158: 117-166. DOI & PDF: 10.1016/j.revpalbo.2009.08.004