Dr David Unwin

Dr David UnwinReader in Palaeobiology

Tel: +44 (0) 116 252 3947

Email: dmu1@le.ac.uk




Personal details

  • Programme Director, Campus-based Museum Studies

My doctoral studies on Cretaceous pterosaur taxonomy and systematics were conducted under the supervision of the late Bev Halstead and I was awarded my PhD by Reading University in 1991.

Following several long term research visits to the USSR between 1988 and 1991, where I was hosted by the Kafedra Palaeontologie in Moscow State University in what was then the USSR, I took up a Royal Society Research Fellowship in the Dept. of Earth Sciences, University of Bristol, in 1992.

During my seven years in Bristol, I developed a broad range of research projects on pterosaurs and other Mesozoic vertebrates, aided by a string of PhD students and several joint programs with colleagues in Russia, China and the US. I was also widely involved in teaching, particularly within palaeontology and helped to establish the highly successful MSc in Palaeobiology.

In 1998 I was appointed Curator of Fossil Reptiles and Birds in the Museum für Naturkunde, Berlin. In addition to caring for the collections, which included the Berlin Archaeopteryx - arguably the most famous fossil in the world - I used my time in Berlin to further develop my research program on pterosaurs, a highlight of which was the publication in 2005, by Pi Press, of The Pterosaurs.

I also helped to develop an application for and subsequently led the German section of the EU funded museums-based project SYNTHESYS, and was the principal scientific officer responsible for the redevelopment of the Dinosaur Hall, which re-opened in 2007.

In 2006, I moved to the School of Museum Studies, Leicester, and in 2009, was appointed as Reader in Palaeobiology.


In my role as Programme Director for Museum Studies. I represent the interests of all campus-based students pursuing a Masters degree (MA or MSc) in Museum Studies.

I am also responsible for:

  • overseeing curriculum development
  • assessment
  • student progress
  • learning resources
  • quality assurance

for this degree course. I also assess all MS applications and serve on the School’s Exhibitions committee and the Senate Student Discipline Committee.


I contribute to several areas of the campus-based Masters degrees including collections care and management and exhibitions.

I am responsible for the Option on Natural History Curatorship, organise the Option module and supervise the Think Tanks.


Codorniú, L., Paulina-Carabajal, A., Pol, D., Unwin, D. M. & Rauhut, O. W. M. A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ (in press).

Unwin, D. M. Hands off, its my collection! Fossils, museums and the problems of access. European Association of Vertebrate Palaeontology (in press).

Unwin, D. M. & Deeming, D. C. The reproductive biology of pterosaurs. European Association of Vertebrate Palaeontology (in press).

Unwin, D. M. 2015. Non-pterodactyloid monofenestratans – rewriting the evolutionary history of pterosaurs. Pp.50-52. In: Witton et al., (eds) Flugsaurier 2015, University of Portsmouth Press.

Unwin, D. M. & Deeming, D. C. 2015. Growth rates and their contraints in pterosaurs. pp.52-54. In: Witton et al., (eds) Flugsaurier 2015, University of Portsmouth Press.

Unwin, D. M., Lü Junchang, Pu Hanyong & Jin Xingsheng. 2014. Pterosaur tails tell tales of modularity and heterochrony in the evolution of the pterodactyloid bauplan. 74th Meeting of the Society of Vertebrate Paleontology (November 5-8, 2014). Berlin (Germany) p.244.

Unwin, D. M., Martill, D. & Hing, R. 2013. DGM 1475-R, a fragment of three-dimensionally preserved pterosaur wing membrane from the Santana Formation of Brazil. Pp. 43-44. In: Walsh, S. et al., (eds). 61st Symposium on Vertebratye Palaeontology and Comprative Anatomy, 74th Meeting of the Society of Vertebrate Paleontology (2013).

Lü Junchang, Unwin, D. M., Zhao Bo, Gao Chunling & Shen Caizhi. 2012. A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae) from the Middle/Upper Jurassic of Qinglong, Hebei Province, China. Zootaxa, 3158: 1-19.

Lü Junchang, Unwin, D. M., Deeming, D. C., Jin Xingsheng, Liu Yongqing and Ji Qiang. 2011. An egg-adult association, gender, and reproduction in pterosaurs. Science, 331, 321-324..

Martill, D. M. & Unwin, D. M. 2011. The world’s largest toothed pterosaur, NHMUK R481, an incomplete rostrum of Coloborhynchus capito (Seeley 1870) from the Cambridge Greensand of England, Cretaceous Research, 34, 1-9.

Remes, K., Unwin, D. M., Klein, N., Heinrich, W.-D. & Hampe, O. 2011. Skeletal reconstruction of Brachiosaurus brancai in the Museum für Naturkunde, Berlin: summarizing 70 years of sauropod research. In: Klein, N., Remes, K. & Sander, P. M. (eds.). Biology of the Sauropod Dinosaurs: Understanding the Life of Giants. Indiana University Press, Bloomington, pp. 305–316.

Ibrahim, N., Unwin D. M., Martill, D. M., Baidder L. & Zouhri, S. 2010. A new pterosaur (Pterodactyloidea: Azhdarchidae) from the Upper Cretaceous of Morocco. PLoS ONE, 5(5): e10875.

Lü Junchang, Unwin, D. M., Ji Shu-an & Ji Qiang (Eds). Flugsaurier 2010, Third International Symposium on Pterosaurs: 5-10 August, 2010, Beijing, China. Acta Geoscientica Sinica, 31 (Supp. 1) 1-93.

Lü Junchang, Unwin, D. M. & Jin Xingsheng. 2010. Modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B: Biological Sciences, 277, 383-389.

Claessens, L. P. A. M, O'Connor, P. M. and Unwin, D. M. 2009. Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism. PLoS ONE, 4(2): e4497.

Unwin, D. M. and Deeming, D. C. 2008. Pterosaur eggshell structure and its implications for pterosaur reproductive biology. Zitteliana, 28, 199-207.

Lü, Junchang, Unwin, D. M., Xu Li and Zhang Xingliao. 2008. A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution. Naturwissenschaften, 95, 891-897.

Obata I., Shibata K., Matsukawa, M. and Unwin, D. M. 2007. New record of a pterosaur from the Late Cretaceous Izumi Group, Awaji Island, Hyogo Prefecture, Japan. Annual Reports of the Fukada Geological Institute, 8, 1-14.

Schaumberg, G., Unwin, D. M. and Brandt, S. 2007. New information on the anatomy of the Late Permian gliding reptile Coelurosauravus. Paläontologisches Zeitschrift, 81/2, 160-173.

Unwin, D. M., Zhou Zhonghe, Yabumoto, Y., Ohashi, T., Ueda, K. and Baba, M. 2007. Pterosaurs: Rulers of the Skies in the Age of Dinosaurs. Asahi Shimbun, Osaka, 92pp.

Unwin, D. M. and Martill, D. M. 2007. Pterosaurs from the Lower Cretaceous Crato Formation of the Chapada do Araripe, north east Brazil. Pp. 475-524. In: Martill, D. M. (ed.), The Fossils Of The Crato Formation: Window On An Ancient World. Cambridge University Press, Cambridge, UK, 625pp. ISBN: 978-0-521-85867-0. [Chapter 17]

Unwin, D. M. 2007. Life takes to the air. Pp. 174-175. In: Glaubrecht, M., Kinitz, A. and Moldrzyk, U. (eds), Evolution in action. Prestel Verlag, Berlin, Germany, 248pp. ISBN: 978-3-7913-3857-6.

Unwin, D. M ., Lü Junchang and Deeming, D. C. 2006. Were all pterosaurs oviparous? Pp. 143-169. In: Papers from the 2005 Heyuan International Dinosaur Symposium. Geological Publishing House, Beijing.

Lü, Jun-chang, Jin, Qinsheng, Unwin, D. M., Zhao, Lijun, Azuma, Y. 2006. A new species of Huaxiapterus (Pterosauria: Pterodactyloidea) from the Lower Cretaceous of Western Liaoning, China with comments on the systematics of tapejarid pterosaurs. Acta Geologica Sinica, 80, 315-326.

Unwin, D. M. 2005. The Pterosaurs from Deep Time. Pi Press, New York, 347pp.

Wilkinson, M. T., Unwin, D. M. & Ellington, C. P. 2005. Lift capability of pterosaur wings and the evolution of gigantism. Proceedings of the Royal Society B: Biological Sciences. 273, 119-126.

Steel, L., Martill, D. M., Unwin, D. M. & Winch, J. D.2005.‘A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England. Cretaceous Research, 26, 686-698.

Tischlinger, H. & Unwin D. M. 2005. Ultra-violet light investigation of the Berlin example of Archaeopteryxlithographica H. v. Meyer 1861 and the isolated Archaeopteryx feather. Archaeopteryx, 23, 17-50.

Deeming, D. C. and Unwin, D. M. 2004. Reptilian incubation: evolution and the fosil record. In Deeming D. C. (ed.), Reptilian Incubation: Environment, Evolution and Behaviour. Nottingham University Press, Nottingham. p. 1-14.

Unwin, D. M. 2004. Eudimorphodon and the early history of pterosaurs. In: Renesto, S. & Pagganoni, A. (eds), A symposium honouring the 30th anniversary of the discovery of Eudimorphodon, Bergamo 2003. Riv. Mus. civ. Sc. Mat. "E. Caffi" Bergamo, 22, 41-48.

Bakhurina, N. N. & Unwin, D. M. 2004. Reconstructing the flight apparatus of Eudimorphodon . In: Renesto, S. & Pagganoni, A. (eds), A symposium honouring the 30th anniversary of the discovery of Eudimorphodon, Bergamo 2003. Riv. Mus. civ. Sc. Mat. "E. Caffi" Bergamo, 22, 7-10.

Unwin, D. M. 2003. Smart-winged pterosaurs. Nature, 425, 910-911.

Unwin, D. M. 2003. On the phylogeny and evolutionary history of pterosaurs. In: Buffetaut, E. & Mazin, J.-M. (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217, 139-190.

Carpenter, K., Unwin, D. M., Cloward, K. & Miles, C. A. 2003. new scaphognathine pterosaur from the Upper Jurassic Morrison Formation, Bone Cabin Quarry, Wyoming, USA. In: Buffetaut, E. & Mazin, J.-M. (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217, 45-54.

Unwin, D. M. 2002. On the systematic relationships of Cearadactylusatrox, an enigmatic Early Cretaceous pterosaur from the Santana Formation of Brazil. Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe, 4, 237-261.

Hwang, Koo-Geun, Huh, M., Lockley, M. G., Unwin, D. M. & Wright, J. L. 2002. New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, SW Korea. Geological Magazine, 139 (4), 421-435.

Unwin, D. M. 2002. Pterosauria (Pterosaurs). In: Encyclopedia of Life Sciences, 15, 700-702. London: Nature Publishing Group.

Unwin, D. M. 2001. An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England. Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe, 4, 189-222.

Unwin, D. M. Patterns and processes of gigantism in pterosaurs. International Congress of Vertebrate Morphology IV, Journal of Morphology, 248, 292. [Abstract].

Unwin, D. M. 2001. Pterosaur locomotion. In: Briggs, D.E.G. & Crowther, P., (eds), Palaeobiology II, 422-425.

Unwin, D. M. & Benton, M.J. 2001. Longisquama fossil and feather morphology. Science. 291, 1899.

Unwin, D. M. & Henderson, D. 2001. Ein Saurier bewegt sich wieder. Theatrum Naturae, 1, 7-9.

Benton , M. J., Kurochkin, E., Shiskin, M. & Unwin, D. M. (eds) 2000. The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, 696pp.

Unwin, D. M. & Bakhurina, N. N. 2000. Pterosaurs from Russia, Middle Asia and Mongolia. In: Benton, M.J., Kurochkin, E., Shiskin, M. and Unwin, D. M. (eds), The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, pp. 420-433.

Unwin, D. M., Alifanov, V. A. & Benton, M. J. 2000. Enigmatic small reptiles from the Late Triassic of Kirghizia. In: Benton, M. J., Kurochkin, E., Shiskin, M. and Unwin, D. M., (eds), The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, pp. 177-186.

Unwin, D. M., Lü, J. & Bakhurina, N. N. 2000. On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China. Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe, 3, 181-206.

Unwin, D. M. & Heinrich, W.-D. 1999. On a pterosaur jaw remain from the Late Jurassic of Tendaguru, East Africa. Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe, 2, 121-134

Unwin, D. M. 1999. Pterosaurs: back to the traditional model? Trends in Evolution and Ecology, 14(7), 263-268.

Unwin, D. M. 1999. Cretaceous fossil vertebrates (Editor). Special Paper in Palaeontology,60, 1-219.

Evans, S. E., Manabe, M., Cook, E., Hirayama, R., Isaji, S., Nicholas, C., Unwin, D. M., & Yabumoto, Y. 1998. An Early Cretaceous assemblage from Gifu Prefecture, Japan. In: Lucas, S. G., Kirkland, J. I. and Estep, J. W. (eds), Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science, Bulletin, 14: 183-186.

Unwin, D. M. 1998. Feathers, filaments and theropod dinosaurs. Nature 391: 119-120. (reprinted in Gee, H. (ed.) 2001, 'Rise of the Dragon', University of Chicago Press, Chicago, 83-86).

Unwin, D. M. 1997. The new dinosaur book. Aladdin Books, London, 32 pp. [Book].

Wright, J. L., Unwin, D. M., Lockley, M. G. & Rainforth, E. 1997. Pterosaur tracks from the Purbeck Formation of Dorset, England. Proceedings of the Geologist's Association, 108: 39-48.

Martill, D.M. & Unwin, D. M. 1997. Small spheres in fossil bones: blood corpuscles or diagenetic artefacts? Palaeontology. 40: 619-624.

Kemp, R. A. & Unwin, D. M. 1997. The taphonomy of Archaeopteryx. Lethaia, 30: 229-238.

Lockley, M.G., Huh, M., Lim, S.-K., Yang, S.-Y., Chun, S.-S., & Unwin, D. M. 1997. First report of pterosaur tracks from Asia, Chullanam Province, Korea. Journal of the Palaeontological Society of Korea, 2: 17-32.

Unwin, D. M. , Bakhurina, N. N., Lockley, M. G., Manabe, M., & Lü, J. 1997. Pterosaurs from Asia. Journal of the Palaeontological Society of Korea,2: 43-65.

Unwin, D. M. & Lü Junchang 1997. On Zhejiangopterus and the relationships of pterodactyloid pterosaurs. Historical Biology, 12: 199-210.

Unwin, D. M. 1997. Pterosaur tracks and the terrestrial ability of pterosaurs. Lethaia, 29: 373-386.

Unwin, D. M. 1996. The fossil record of Middle Jurassic pterosaurs. In: Morales, M. (ed.), The Continental Jurassic, Museum of Northern Arizona Bulletin, 60: 291-304.

Unwin, D. M. 1996. Mysteries of prehistoric life. Aladdin, London. 40 pp. [Book].

Unwin, D. M. , Hasegawa, Y., Shimizu, K. & Manabe, M. 1996. First record of pterosaurs from the Early Cretaceous Tetori Group: a wing-phalange from the Amagodani Formation in Shokawa, Gifu Prefecture, Japan. Bulletin of the National Science Museum, Tokyo, Series C (Geology and Paleontology), 22, 37-46.

Unwin, D. M. , Frey, E., Martill, D. M., Clarke, J. & Riess, J. 1996. On the nature of the pteroid in pterosaurs. Proceedings of the Royal Society: Biological Sciences, 263, 45-52.

Bakhurina, N. N. & Unwin, D. M. 1995. A survey of pterosaurs from the Jurassic and Cretaceous of the former Soviet Union and Mongolia. Historical Biology, 10, 197-245.

Bakhurina, N. N. & Unwin, D. M. 1995. A preliminary report on the evidence for 'hair' in Sordes pilosus, an Upper Jurassic pterosaur from Middle Asia. In: Sun, A. and Wang Y. (eds) Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota. Short Papers, pp. 79-82.

Unwin, D. M. 1995. Preliminary results of a phylogenetic analysis of the Pterosauria (Diapsida: Archosauria). In: Sun, A. and Wang Y. (eds) Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota. Short Papers, pp. 69-72.

Unwin, D. M. & Bakhurina, N. N. 1995. Wing shape in pterosaurs. Nature, 374, 316.

Unwin, D. M. & Bakhurina, N. N. 1994. Sordes pilosus and the nature of the pterosaur flight apparatus. Nature, 371, 62-64.

Unwin, D. M. 1994. Brachiosaurus. Kingfisher, London, 39 pp. [Book].

Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.), The Fossil Record 2, pp 717-737, Chapman and Hall, London.

Deeming, D. C., Halstead, L. B., Manabe, M. Unwin, D. M. 1993. A true ichthyosaur embryo with comments on ichthyosaur ontogeny. Modern Geology, L. B. Halstead Memorial Volume, 18 (4), 423-442.

Unwin, D. M. 1992. Wings over the dinosaur world. Wildlife Fact File Yearbook1993: 54-59.

Unwin, D. M. 1991. The morphology systematics and evolutionary history of pterosaursfrom the Cretaceous Cambridge Greensand of England. PhD thesis, Department of Zoology, Reading University, 527 pp.

Martill, D. M., Wilby, P. & Unwin, D. M. 1990. Stripes on a pterosaur wing. Nature, 346: 116.

Fraser , N.C. & Unwin, D. M. 1990. Pterosaur remains from the Upper Triassic of Britain. Neues Jahrbuchfur Geologie und Palaeontologie Monatsheft, 1990(5): 272-282.

Unwin, D. M. 1989. A predictive method for the identification of vertebrate ichnites and its application to pterosaur tracks. In: Gillette, D. M. and Lockley, M. (eds), Dinosaur tracks and traces, pp 259-274, Cambridge University Press, Cambridge.

Martill, D. M. & Unwin, D. M. 1989. Exceptionally well preserved pterosaur wing membrane from the Cretaceous of Brazil. Nature, 340: 138-139.

Unwin, D. M. 1988. New pterosaurs from Brazil. Nature, 332: 398-399.

Unwin, D. M. 1988. A new pterosaur from the Kimmeridge Clay of Kimmeridge, Dorset. Proceedings of the Dorset Natural History and Archaeological Society, 109: 150-153.

Unwin, D. M. 1988. Extinction and survival in birds. In: Larwood, G.P. (ed.), Extinction and Survival in the Fossil Record. Systematics Association Special Volume 34, 295-318, Clarendon Press, Oxford.

Unwin, D. M. 1988. New remains of the pterosaur Dimorphodon (Pterosauria: Rhamphorhynchoidea) and the terrestrial ability of early pterosaurs. Modern Geology, 13: 57-68.

Unwin, D. M. 1987. Pterosaur locomotion. Joggers or waddlers? Nature, 327: 13-14.

Unwin, D. M. 1987. Pterosaur extinction: nature and causes. Memoires Sociéte Geologique de France, 150: 105-111.

Unwin, D. M. 1986. Electric pterosaur takes off. Geology Today, 2(1): 17.


I am broadly interested in all fossil vertebrates, but have chosen to focus my research effort on a single group: pterosaurs, extinct flying reptiles also known as pterodactyls.

During the last 30 years I have published on many different aspects of pterosaurs including their:

  • anatomy
  • flight
  • walking abilities
  • tracks
  • respiration
  • reproductive biology
  • growth
  • origins
  • interrelationships
  • evolutionary history

By integrating the results of these studies with the large and rapidly expanding body of research on pterosaurs, my aim is to develop a more comprehensive and consistent understanding of the palaeobiology of these extraordinary animals.

Since arriving at Leicester in 2006, much of my research has focused on projects involving the taxonomy and phylogeny of pterosaurs. The principal achievement here has been the discovery and description of Darwinopterus, a pterosaur from the Jurassic of China, that my colleagues and I named after Charles Darwin on the occasion of the 150th anniversary of the publication of ‘On the origin of species’.

This remarkable pterosaur has an almost perfect ‘transitionary’ anatomy that shows, for the first time, how long-tailed basal pterosaurs evolved into short tailed, relatively advanced pterodactyloid pterosaurs. It also points to an unusual ‘modular’ mode of evolution that helps to explain some surprising patterns in the fossil record such as evidence for rapid, large-scale changes.

In addition to Darwinopterus, I have also helped to describe several new pterosaurs including Shenzhoupterus and Qinglongopterus from China and Alanqa from Africa, and am currently involved in co-editing the first comprehensive volume on pterosaur systematics, The Pterosauria with Dave Martill (University of Portsmouth).

Contemporaneous with systematic studies, I have also been leading a series of related projects on the reproductive biology and growth of pterosaurs funded by a Royal Society International Joint Project Grant and involving colleagues from China and the UK. A highlight of this work has been the discovery of a female example of Darwinopterus, ‘Mrs T’, preserved in association with an egg. This spectacular find demonstrates that the showy head crests borne by many male pterosaurs were used for display, either to attract a mate or intimidate rivals. ‘Mrs T’ also shows that pterosaurs laid relatively small eggs with pliable shells. So, as in some living reptiles, pterosaur eggs were likely incubated by burial in vegetation or soil, rather than brooded as happens in birds.

In 2009, I was invited to contribute to a 3D film project on pterosaurs hosted by David Attenborough and produced by Sky Atlantic. Following filming in 2010, ‘Flying Monsters’ was released across an array of media platforms including Imax cinemas, TV, DVD and on the web. Content can also be viewed via an app for the iPad and Android.

The film, which has already generated more than $10 million in revenue, met with critical acclaim and has won a string of prizes including a BAFTA in 2011 for the best specialist factual film. ‘Flying Monsters’ also achieved impact in other ways, acting as a ‘showcase for 3D moviemaking as applied to documentaries’ and providing content for educational websites such as that hosted by the National Geographic.


I second supervise 10 PhD students.

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