Publications

BARYONYCHINE BRAINS

We CT-scanned the braincases of the British spinosaurid dinosaurs Baryonyx walkeri from Surrey and Ceratosuchops inferodios from the Isle of Wight and here present digital endocasts of brain anatomy. Overall, the brains are morphologically similar to those of other non-coelurosaurian tetanurans. The paper is OA…

The Isle of Wight baryonychine spinosaurid Ceratosuchops interacts with another animal, with the position and anatomy of its brain (and associated structures) shown in the inset. Image: Anthony Hutchings.

Barker, C. T., Naish, D., Trend, J., Michels, L. V., Witmer, L., Ridgley, R., Rankin, K., Clarkin, C. E., Schneider, L. & Gostling, N. J. 2023. Modified skulls but conservative brains? The palaeoneurology and endocranial anatomy of baryonychine dinosaurs (Theropoda: Spinosauridae). Journal of Anatomy doi: 10.1111/joa.13837

• A Tetrapod Zoology write-up of this paper is here… A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains

CRYPTOZOOLOGY AS CULTURE, AGAIN

Titled ‘A Cultural Phenomenon’, this article discusses my (by now very familiar) argument (espoused in my 2017 book Hunting Monsters and elsewhere) that many so-called cryptids are products of culture and lore, not new animal species awaiting discovery (Naish 2022: digital version here). Nevertheless, discussions of these alleged creatures are so entwined with writings on zoology and natural history that they are very much part of zoological history, real or not. A slightly controversial claim made in the article is that cryptozoology as currently conceived is creeping away from the zoology-based version of the subject favoured by Bernard Heuvelmans and his associates, and is instead becoming a vague, pseudoscientific topic where people think that bigfoot and Nessie should be discussed alongside demons, floating lights, UFOs and portals to other dimensions. There’s evidence that this is indeed the case within the popular sphere, but the pushback view is that ‘the popular sphere’ doesn’t represent the views and actions of authors and researchers.

This article made the cover. It includes images of cryptids provided by Tyler Stone.

Naish, D. 2022. A cultural phenomenon. The Biologist 69 (3), 16-21.

THE EOTYRANNUS MONOGRAPH

At long last, the monographic description of the Wessex Formation tyrannosauroid Eotyrannus lengi Hutt et al., 2001 is published. This work substantially updates the preliminary 2001 Cretaceous Research paper (available below) and provides a wealth of new data on this dinosaur’s anatomy. We also include a new analysis of Tyrannosauroidea and provide a new placement for Eotyrannus (and other tyrannosauroid taxa), feature a new skeletal reconstruction, and discuss how it compares to other described Wealden theropod taxa. This work formed the bulk of my PhD thesis (completed in 2006) and finally seeing it in print is a significant personal achievement.

Naish, D. & Cau, A. 2022. The osteology and affinities of Eotyrannus lengi, a tyrannosauroid theropod from the Wealden Supergroup of southern England. PeerJ 10:e12727.

• A Tetrapod Zoology write-up of this paper is here… The Eotyrannus Monograph Is Finally Published

ANOTHER WEALDEN SPINOSAURID: THE ‘WHITE ROCK SPINOSAURID’

Several new bones from the Lower Cretaceous Vectis Formation of the Isle of Wight, discovered by local collectors Nick Chase and Mark Penn and Wealden dinosaur specialist Jeremy Lockwood, prove to belong to a giant spinosaurid. This is the first member of the group from the Vectis Formation and augments our poor knowledge of the Vectis Formation’s dinosaur assemblage. Anatomical details and phylogenetic analysis suggest that this animal (a new taxon, but one we opted not to name) is perhaps a spinosaurine. It is also gigantic: the dimensions of the elements suggest a length exceeding 10 m, and perhaps a size exceeding that of Spinosaurus.

Composite showing the best of the White Rock spinosaurid bones and where we think they come from. Silhouette by Dan Folkes.

Barker, C. T., Lockwood, J. A. F., Naish, D., Brown, S., Hart, A., Tulloch, E. & Gostling, N. J. 2022. A European giant: a large spinosaurid (Dinosauria: Theropoda) from the Vectis Formation (Wealden Group, Early Cretaceous), UK. PeerJ 10:e13543.

• A Tetrapod Zoology write-up of this paper is here… A Giant Spinosaurid Dinosaur from the Cretaceous of the Isle of Wight

TWO NEW ISLE OF WIGHT SPINOSAURID THEROPODS

Between 2013 and 2017, a large number of bones belonging to two Baryonyx-like theropods were discovered in the Wessex Formation at Chilton Chine, Isle of Wight. Both specimens proved to differ from Baryonyx walkeri in numerous detailed respects and, surprisingly, grouped with the African Suchomimus phylogenetically. In a new study led by Chris Barker and Neil Gostling at the University of Southampton, we analysed these two new animals and named them: they are Ceratosuchops inferodios and Riparovenator milnerae. This paper is a preliminary report and a more detailed analysis will be published later.

Life reconstruction of Ceratosuchops (at left) and Riparovenator, shown sharing a flooded depression on the Wessex Formation floodplain. Image: Anthony Hutchings.

Barker, C. T., Hone, D. W. E., Naish, D., Cau, A., Lockwood, J. A. F., Forster, B., Clarkin, C. E., Schneider, P. & Gostling, N. J. 2021. New spinosaurs from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports 11: 19340.

• A Tetrapod Zoology write-up of this paper is here… Two New Spinosaurid Dinosaurs from the English Cretaceous

ENOUGH WITH ‘CETARTIODACTYLA’ ALREADY

The idea that whales (cetaceans) are deeply nested within Artiodactyla is now well-supported by molecular, fossil and anatomical data. Cetacea is thus part of Artiodactyla. Some molecular biologists have wrongly thought that this discovery warrants a renaming of Artiodactyla, and thus we have the redundant ‘Cetartiodactyla’. This brief paper explains why the name is not needed and should be dropped. It is only one of several examples of this sort of thing: others include ‘Euarchonta’ and ‘Eulipotyphla’.

Prothero, D. R., Domning, D., Fordyce, R. E., Foss, S., Janis, C., Lucas, S., Marriott, K. L., Metais, G., Naish, D., Padian, K., Rössner, G., Solounias, N., Spaulding, M., Stucky, R. M., Theodor, J. & Uhen, M. 2021. On the unnecessary and misleading taxon “Cetartiodactyla”. Journal of Mammalian Evolution http://doi.org/10.1007/s10914-021-09572-7

The consensus topology of Artiodactyla, showing how whales are nested within artiodactyls, not a sister-taxon to that order (Original drawings by K. L. Mariott).

THE DINOSAUROID REVISITED, ALMOST 40 YEARS ON

In 1982, palaeontologist Dale Russell and model-maker and taxidermist Ron Séguin proposed that troodontid theropods might have evolved into humanoids – ‘dinosauroids’ – had they not gone extinct. Dale Russell’s death in 2019 resulted in the compilation of a special Russell-themed issue of Canadian Journal of Earth Sciences. Thanks to special access to Russell’s correspondence, and new info provided by Ron Séguin, I and Will Tattersdill have produced a new review of the dinosauroid’s backstory and impact. We cover several topics not discussed before, including Russell’s correspondence with Carl Sagan, his involvement with SETI, and the possible relevance of other fictional humanoid reptiles. A section on the possible impact of Russell’s religious views and his preference for Pierre Teilhard de Chardin’s argument that the evolution of humanoids was inevitable (and that humanoids have a special place in the design of the universe) was deleted from the paper by the editors.

Naish, D. & Tattersdill, W. 2021. Art, anatomy and the stars: Russell and Séguin’s dinosauroid. Canadian Journal of Earth Sciences https://doi.org/10.1139/cjes-2020-0172

• A Tetrapod Zoology write-up on this paper is here… Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old

NEWLY HATCHED PTEROSAURS WERE CAPABLE FLIERS

The idea that hatchling pterosaurs – termed ‘flaplings’ in the writings of pterosaur expert David Unwin – might have been precocial and able to fly immediately is not new. We aimed to test the idea further by examining the aerodynamic abilities of a few specific juvenile pterosaur specimens: our inclusion in particular of the ‘Nemicolopterus crypticus‘ specimen is important since this animal was described as a distinct taxon but is argued here to be a juvenile Sinopterus. If pterosaur hatchlings were as flight-capable as we argue, they very likely lived different lives from their parents and occupied very different ecological niches… (art below by Mark Witton)

Naish, D., Witton, M. P. & Martin-Silverstone, E. 2021. Powered flight in hatchling pterosaurs: evidence from wing form and bone strength. Scientific Reports 11:13130.

• A Tetrapod Zoology write-up on this paper is here… Baby Pterosaurs Were Excellent Fliers and Occupied Different Niches From Their Parents

ON BEING A SCIENCE WRITER

Everyone who breaks into the world of professional writing gets asked – a lot – how they did it. The problem is that everyone’s story is different and that there’s no one way. Furthermore, I think it’s important to be honest and say that making a living from writing is really, really hard and not something I’d recommend. In this article, I explained my own adventures in science writing and my thoughts on how writing should perhaps be done.

Naish, D. 2021. Becoming a science writer. Biological Sciences Review 33 (3), 27-29. [pdf here]

ICHTHYOSAURS OF THE KIMMERIDGE CLAY

In this review article, we look at the several ichthyosaur taxa from the Middle Jurassic Kimmeridge Clay Formation. The article was intended to see print prior to the publication of several reviews and revisions of Kimmeridge Clay ichthyosaurs produced by other authors but, unfortunately, appeared in print long after those other works… the result being that it was substantially out of date even at the time of its publication, oh well. We promote the synonymisation of Grendelius with Brachypterygius (whereas more recent work indicates that the two should be kept separate), described Nannopterygius as being known from only one good specimen (whereas more recent work reveals that there are many), and were unable to incorporate the publication of Thalassodraco etchesi Jacobs & Martill, 2020.

Naish, D. & Moon, B. 2020. Ichthyosaurs. In Martill, D. M. & Etches, S. (eds). Fossils of the Kimmeridge Clay Formation 2. The Palaeontological Association, London, pp. 75-90.

At right, the holotype Grendelius mordax skull (CAMSM J68516) on display at the Sedgwick Museum of Earth Science, University of Cambridge, UK. At left, an accompanying model depicting the animal in life. Images: Darren Naish.

• A Tetrapod Zoology article on this… article is here: The Ichthyosaurs of the Kimmeridge Clay

‘UNEXPECTED ISLE OF WIGHT AIR-FILLED HUNTER’, A NEW THEROPOD

A series of theropod dinosaur vertebrae were discovered at Knock Cliff, Shanklin, on the south-east coast of the Isle of Wight, and originated from the Ferruginous Sandstone Formation of the Greensand Group of the Isle of Wight. The vertebrae impressed us in being highly distinctive anatomically, and in being especially pneumatic. They clearly represent a new taxon, which we named Vectaerovenator inopinatus Barker et al., 2020. The affinities of Vectaerovenator proved difficult to pin down but it is certainly a tetanuran, and perhaps a coelurosaur and even a tyrannosauroid. The taxon is significant in coming from a poorly sampled section of the Cretaceous from which few European theropod remains are known. We hope that future finds will improve our knowledge of this intriguing animal.

Barker, C. T., Naish, D., Clarkin, C. E., Farrell, P., Hullmann, G., Lockyer, J., Schneider, P., Ward, R. K. C. & Gostling, N. J. 2020. A highly pneumatic middle Cretaceous theropod from the British Lower Greensand. Papers in Palaeontology 2020, 1-19.

The corpse of a deceased Vectaerovenator floats in the shallow Early Cretaceous sea covering what’s now southern England. Ammonites and platypterygiine ichthyosaurs are nearby. Image (c) Trudie Wilson

Silhouette of Vectaerovenator showing the approximate position of the elements described in our paper. I deliberately made the animal look like a generic tetanuran theropod that isn’t referable to any specific lineage. Image: Barker et al. (2020).

• A Tetrapod Zoology discussion of this new taxon and its implications is here: Introducing ‘Unexpected Isle of Wight Air-Filled Hunter’, a New English Theropod Dinosaur

THEROPOD PHYLONYMS CHAPTER

The long-awaited and gigantic Phylonyms volume appeared in print in 2020. I and colleagues authored the theropod section. I regret I do not have a pdf.

Naish, D., Cau, A., Holtz, T. R. Jr, Fabbri, M. & Gauthier, J. A. 2020. Theropoda O.C. Marsh 1881 [D. Naish, A. Cau, T.R. Holtz, Jr., M. Fabbri, and J.A. Gauthier], converted clade name. In de Queiroz, K., Cantino. P. D. & Gauthier, J. A. (eds) Phylonyms: A Companion to the PhyloCode. CRC Press, pp. 1235-1246.

WAS PHOTOLUMINESCENCE PRESENT IN DINOSAURS AND PTEROSAURS?

The discovery of photoluminescent tissues in modern birds (like puffins and rhinoceros auklets), lizards, amphibians and other vertebrates led myself, Cary Woodruff and Jamie Dunning to discuss the possibility that Mesozoic archosaurs might similarly have emitted photoluminescence and perhaps been able to detect it. If the extravagant structures of these animals evolved as signalling structures, could this phenomenon enhance their signalling function? Our paper is a speculative discussion.

Woodruff, D. C., Naish, D. & Dunning, J. 2020. Photoluminescent visual displays: an additional function of integumentary structures in extinct archosaurs? Historical Biology DOI: 10.1080/08912963.2020.1731806 [pdf here]

Would a live dinosaur – like this heterodontosaur – look utterly different if its tissues were photoluminescent? Brian Engh explored this possibility in this excellent piece of art, included in Woodruff et al. (2020). Image: Brian Engh.

• A Tetrapod Zoology write-up of this research is here:  Did Dinosaurs and Pterosaurs ‘Glow’? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays (January 2019)
• Media coverage here: Did dinosaurs glow in the dark? Scientists suggest photoluminescent skin (MetroUK, March 2020)

SEA MONSTERS AND THE ‘PLESIOSAUR EFFECT’

The ‘sea monsters’ reported by witnesses are a diverse lot, and how they’ve been described has changed to some degree over the decades and centuries. Charles Paxton and I have a serious interest in analysing cryptozoological data and publishing those results in the technical literature. We wondered whether the 1800s discovery of fossil plesiosaurs influenced those people who claimed to have seen sea monsters.

Paxton, C. G. M. & Naish, D. 2019. Did nineteenth century marine vertebrate fossil discoveries influence sea serpent reports? Earth Sciences History 38, 16-27. [pdf here]

Some of our graphs (Paxton & Naish 2019). Eyewitnesses increasingly mentioned ‘plesiosaurs’ and the presence of necks throughout the 1800s. Across the same time frame, references to serpent-like features were in decline. Image: Paxton & Naish (2019).

• A Tetrapod Zoology write-up of this research is here: Sea Monster Sightings and the ‘Plesiosaur Effect’ (April 2019)
• Media coverage here: Ancient marine reptile fossil findings influenced sea serpent sightings, new research reveals (History Scotland, April 2019)

A MIXED-SPECIES NESTING ASSEMBLAGE IN LATE CRETACEOUS ROMANIA

Our initial paper on an eggshell-dominated sediment lens from the Maastrichtian of Romania (Vremir et al. 2012) missed the fact that some fossil fragments included in the assemblage demonstrate the presence of a few species in addition to the enantiornithine which dominates it. In this new study, we use SEM microscopy to demonstrate the presence of an additional bird taxon as well as a gekkotan lizard and a crocodylomorph. The site perhaps represents a multi-species assemblage and may show that several species were nesting in close proximity, and perhaps even sharing the same immediate nesting environment.

Fernández, M. S., Wang, X., Vremir, M., Lauren, C., Naish, D., Kaiser, G. & Dyke, G. 2019. A mixed vertebrate eggshell assemblage from the Transylvanian Late Cretaceous. Scientific Reports 9, 1944.

‘Geckoid eggshell’ in the Oarda de Jos assemblage, as seen via SEM. The images (note the different scales) show (A) a distinct two-layered structure and (B) numerous tiny holes in the prisms of the second later. Image: Fernández et al. (2019).

• Tetrapod Zoology write-up of this research is here: A Multi-Species Nesting Assemblage in the Late Cretaceous of Europe (February 2019)

PTEROSAUR PALAEONEUROLOGY

The British azhdarchoid pterosaur Vectidraco (published by Naish et al. 2013) was subjected to CT-scanning by Liz Martin-Silverstone. One discovery made during this research was that the lateral excavations on the sides of the vertebrae were not pneumatic openings (as suggested in Naish et al. 2013) but intervertebral foramina (IVF) associated with the spinal nerves. We became interested in discussing the IVF of pterosaurs more generally (they are often figured but never discussed). The neural canal size of Vectidraco was also interesting in that it appeared proportionally larger than that of the other pterosaurs we studied. Does neural canal size have any correlation with locomotor abilities and lifestyle in pterosaurs? Our results suggest that Vectidraco was more proficient at walking than the other pterosaurs we looked at, a result consistent with my previous work on pterosaurs (Witton & Naish 2008). This is a preliminary study involving a small data set. We hope that it inspires additional work on pterosaur palaeoneurology.

Martin-Silverstone, E., Sykes, D. & Naish, D. 2018. Does postcranial palaeoneurology provide insight into pterosaur behaviour and lifestyle? New data from the azhdarchoid Vectidraco and the ornithocheirids Coloborhynchus and Anhanguera. Palaeontology 2018, 1-14. doi: 10.1111/pala/12390

Neural canal cross-sectional area in our three pterosaur taxa: when normalised for centrum size, Vectidraco has a proportionally large neural canal. This composite image incorporates figures from Martin-Silverstone et al. (2018) but was produced by the Palaeontological Association. Image: Martin-Silverstone et al. (2018).

• Tetrapod Zoology write-up of this research is here: Postcranial Palaeoneurology and the Lifestyles of Pterosaurs (August 2018)

‘SENSITIVE SNOUTS’ IN BIG THEROPODS?

We CT-scanned skull bones of the Early Cretaceous allosauroid theropod Neovenator in order to examine its internal structure; a project led by Chris Barker at the University of Southampton. We discovered a complex arrangement of connected channels, ducts and openings which we interpreted as a hypertrophied neurovascular system, the presence of which suggests exceptional facial sensitivity. Our conclusions have since been challenged, since the structures we identified may have housed larger blood vessels and pneumatic structures than we accounted for.

Barker, C. T., Naish, D., Newham, E., Katsamenis, O. L. & Dyke, G. 2017. Complex neuroanatomy in the rostrum of the Isle of Wight theropod Neovenator salerii. Scientific Reports 7, 3749.

• Tetrapod Zoology write-up of this research is here: The Sensitive Face of a Big Predatory Dinosaur (June 2017)
• Guardian piece: Dinosaurs’ sensitive snouts enabled courtship ‘face stroking’, study suggests (June 2017)
• Palaeocast: Neovenator Neurovasculature and the Purpose of Sensitive Snouts (June 2017)
• Earth Archives: Meat-eating dinosaurs had sensitive snouts (June 2017)

PLESIOSAUR SWIMMING: LASERS AND ROBOTS

The four-flipper swimming of plesiosaurs – used consistently by this group of animals for over 150 million years – has long been the source of argument and interpretation. Luke Muscutt’s PhD work at the University of Southampton involved the construction of realistic 3D plesiosaur paddles and their attachment to a robot, the performance of which was analysed in a flume tank. Coloured dyes revealed the pattern of flow over the moving paddles, lasers were used to analyse the specifics of their movement. This initial study involved paddles based on those of an Early Jurassic rhomaleosaurian plesiosaur; subsequent studies will involve data from additional plesiosaur lineages.

Muscutt, L. E., Dyke, G., Weymouth, G. D., Naish, D., Palmer, C. & Ganapathisubramani, B. 2017. The four-flipper swimming method of plesiosaurs enabled efficient and effective locomotion. Proceedings of the Royal Society B 284, 20170951.

Our replica plesiosaur flippers in action, the red and blue dyes aiding visualisation of the vortices and wakes being shed from the flippers. Credit: Luke Muscutt/Muscutt et al. (2017).

• A Tetrapod Zoology write-up of this research is here: The Unique and efficient 4-Flipper Locomotion of Plesiosaurs (August 2017)
• A video on Luke’s research is here… The Hydrodynamics of Plesiosaurs

SOME AZHDARCHIDS WERE THICK-NECKED PREDATORS

The view promoted in Witton & Naish (2008) and Witton & Naish (2015) – that azhdarchids were essentially all alike in proportions and hence similar in lifestyle – is shown to be incorrect via our analysis of cervical vertebrae belonging to the giant Late Cretaceous Romanian azhdarchid Hatzegopteryx. We show that the neck of this animal was broad and thick relative to that of other giant azhdarchids (like Arambourgiania) and mechanically able to resist substantial loads. This view is consistent with the absence of large predators (like theropods) from the same region and suggests that Hatzegopteryx was a predator of animals that could have weighed tens of kilos. This study is consistent with our previous proposal that some azhdarchids were relatively short-necked.

Naish, D. & Witton, M. P. 2017. Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators. PeerJ 5:e2908.

Here’s how EME 315 matches up to other azhdarchid cervical vertebrae as goes length to width ratios – it’s clearly an outlier. We can say similar things for the extremely elongate cervical vertebra of Arambourgiania, shown at the top. Credit: Naish & Witton 2017.

Skeletal reconstructions of the giant azhdarchids (A) Hatzegopteryx and (C) Arambourgiania to scale, showing the markedly different body shapes seemingly present in these animals. (B) shows how broad the back of the skull and neck was in Hatzegopteryx. Contrast this with D-E: the small Quetzalcoatlus species (since named Q. lawsoni), with its 4.6 m wingspan and long, slender neck. Scale bar = 1 m. Credit: Naish & Witton 2017.

• A Tetrapod Zoology write-up of this research is here: Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators (February 2017)

MORE SUPPORT FOR ‘TERRESTRIAL STALKING’ IN AZHDARCHIDS

The view that azhdarchid pterosaurs were ‘terrestrial stalkers’, proposed by Witton & Naish (2008), was challenged in 2013 by Alexander Averianov who argued that the depositional settings of azhdarchid fossils were inconsistent with our proposal, that big theropods made our proposal problematic, and that azhdarchids were more likely ‘scoop-netters’. In our response, we re-examined the environmental context in which azhdarchids have been discovered and showed that the evidence was consistent with their being animals of continental settings. We also looked at the behaviour and ecology of those theropod contemporaneous with azhdarchids to see if they would really present the problem which Averianov argued they would, and examined his ‘scoop-netting’ idea in order to test its viability.

Witton, M. P. & Naish, D. 2015. Azhdarchid pterosaurs: water-trawling pelican mimics or “terrestrial stalkers”? Acta Palaeontologica Polonica 60, 651-660.

A giant azhdarchid is a formidable beast. This illustration – from Witton & Naish (2013) – shows (A) Tyrannosaurus, (B) Romanian maniraptoran Balaur, (C) the long-necked giant azhdarchid Arambourgiania, and (D) a human, all to scale. Image by Mark Witton, from Witton & Naish (2013).

The terrestrial stalking hypothesis is based on multiple lines of evidence: it does not (like so many other hypotheses about pterosaur behaviour and ecology) rely on cherry-picking one or two anatomical or palaeoenvironmental features. This image – from Witton & Naish (2013) – shows how several independent pieces of data all provide support for our hypothesis.

• A Tetrapod Zoology write-up of this research is here: Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were (December 2013)
• Mark Witton’s article: Azhdarchid pterosaurs: ‘terrestrial stalkers’ or pelican-esque ‘scoop-feeders’? (October 2013)

BALAUR APPEARS TO BE AN AVIALAN, NOT A DROMAEOSAURID

The remarkable Late Cretaceous Romanian maniraptoran Balaur bondoc was published as a velociraptorine dromaeosaur. However, several of the features which make it unusual (including a reduced hand skeleton, bowed-out pubic bones and a proportionally long hallux) are more typical of Avialae (the Archaeopteryx + bird lineage) than Dromaeosauridae. While in Romania and with direct access to the specimen, I noticed these features, as did theropod worker and phylogeneticist Tom Brougham. We opted to team up with Andrea Cau, who had also noticed the avialan-like features of Balaur and drawn attention to them. Our phylogenetic analysis of Maniraptora confirmed an avialan placement for this taxon, in which case it has to be identified as a secondarily flightless, large descendant of jeholornithid-grade taxa – an exciting idea. It should be noted that this doesn’t affect the imagined life appearance of Balaur, though those reconstructions which make it look especially ‘Velociraptor-like’ are inconsistent with this proposal. Some other studies have also recovered an avialan position for Balaur. Debate about this taxon’s position will continue, at least until good cranial material is found. Balaur‘s finder – Mátyás Vremir – died in 2020, reducing the odds that additional material will be found any time soon.

Cau, A., Brougham, T. & Naish, D. 2015. The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc (Dinosauria, Maniraptora): dromaeosaurid or flightless bird? PeerJ 3:e1032.

Speculative skeletal reconstruction for Balaur bondoc, showing known elements in white and unknown elements in grey. Note that the integument would presumably have substantially altered the outline of the animal in life. Produced by Jaime Headden, used with permission.

A Balaur bondoc pair in their forested Romanian home, 67 or so million years ago. Artwork by Emily Willoughby.

• A Tetrapod Zoology write-up of this research is here: The Romanian Dinosaur Balaur Seems to Be a Flightless Bird (June 2015)

AN ICZN APPEAL REGARDING TAXONOMIC VANDALISM

Rhodin, A. G. J., Kaiser, H., van Dijk, P. P., Wüster, W., O’Shea, M., Archer, M., Auliya, M., Boitani, L., Bour, R., Clausnitzer, V., Contreras-MacBeath, T., Crother, B. I., Daza, J. M., Driscoll, C. A., Flores-Villela, O., Frazier, J., Fritz, U., Gardner, A., Gascon, C., Georges, A., Glaw, F., Grazziotin, F. G., Groves, C. P., Haszprunar, G., Havaš, P., Hero, J. M., Hoffmann, M., Hoogmoed, M. S., Horne, B. D., Iverson, J. B., Jäch, M., Jenkins, C. L., Jenkins, R. K. B., Kiester, A. R., Keogh, J. S., Lacher Jr., T. E., Lovich, J. E., Luiselli, L., Mahler, D. L., Mallon, D., Mast, R., Mcdiarmid, R. W., Measey, J., Mittermeier, R. A., Molur, S., Mossbrugger, V., Murphy, R., Naish, D., Niekisch, M., Ota, J., Parham, J. F., Parr, M. J., Pilcher, N. J., Pine, R. H., Rylands, A. B., Sanderson, J. G., Savage, J., Schleip, W., Scrocchi, G. J., Shaffer, H. B., Smith, E. N., Sprackland, R., Stuart, S. N., Vetter, H., Vitt, L. J., Waller, T., Webb, G., Wilson, E. O., Zaher, H. & Thomson, S. 2015. Comment on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the availability of the generic name and for the nomenclatural validation of the journal in which it was published. (Case 3601; see BZN 70: 234–237; 71: 30–38, 133–135, 181–182, 252–253). Bulletin of Zoological Nomenclature 72 (1): 65-78. [pdf here]

SOME AZHDARCHIDS HAD ‘SHORT’, ROBUST NECKS

The idea that azhdarchid pterosaurs were anatomically homogenous is challenged by several (mostly poorly known) taxa. These indicate that neck length and proportions, and body size overall, was variable in the group (an idea first mooted by Dave Unwin in his 2006 book The Pterosaurs From Deep Time). LPB (FGGUB) R.2395 is a Haţeg Basin cervical vertebra from a mid-sized azhdarchid, estimated to have a wingspan of 3-4 m. Despite its small size (89 mm long), its smooth (non-pitted), polished external bone texture shows that it belonged to an adult. It is probably a cervical IV. The most interesting thing about R.2395 is that it is relatively broad for its length, this indicating that the animal had a neck length c 30-40% shorter than that of other azhdarchids of similar size. It indicates the presence of an additional azhdarchid taxon in the Haţeg Basin fauna. We also argued in this paper that efforts to sink all Haţeg Basin azhdarchids into a single taxon do not withstand scrutiny. Due to an editorial mishap, the final published version of this paper is not as richly illustrated as planned in earlier drafts.

Vremir, M., Witton, M., Naish, D., Dyke, G., Brusatte, S. L., Norell, M. & Totoianu, R. 2015. A medium-sized robust-necked azhdarchid pterosaur (Pterodactyloidea: Azhdarchidae) from the Maastrichtian of Pui (Haţeg Basin, Transylvania, Romania). American Museum Novitates 3827, 1-16. [pdf here]

At left, R.2395 as reconstructed by Mark Witton: a mid-sized, robust-necked azhdarchid. At right, R.2395 shown in various views.

STATE OF THE PALAEOART

In this essay, Mark, John and I review the history of palaeoart and emphasise its importance in the dissemination of palaeontological knowledge. Given its importance, we decry the lazy approach taken by many concerning plagiarism, poor standards and a ‘culture of copying’; indeed, we were inspired to write the piece following several (then new) cases of such. Part of the problem remains that working palaeontologists are typically expected by publishers and the media to be arbiters or experts on matters concerning palaeoart when in fact they are often (albeit not always) neither interested nor knowledgeable on matters concerned.

Witton, M. P., Naish, D. & Conway, J. 2014. State of the Palaeoart. Palaeontologia Electronica 17, Issue 3; 5E: 10p.

Palaeoart memes and the culture of copying, evidenced by different iterations of Robert Bakker’s 1971 ‘giraffoid’ Barosaurus, a distinctive reconstruction characterised by a mast-like neck, ventral midline ridge on the neck, and (via foreshortening in the original composition) a short tail.

THALASSODROMEUS IS NOT PRESENT IN THE ROMANIAN CRETACEOUS

In 2014, Grellet-Tinner and Codrea made the surprising claim that the azhdarchoid pterosaur Thalassodromeus – known from good remains described from the Aptian of Brazil – had a presence in the Maastrichtian Sebeş Formation of Romania too. The fossil concerned (an irregularly shaped sheet of bone) was identified as the premaxillary crest of a new species (T. sebesensis) and suggested to have implications for pterosaur diversity in the Late Cretaceous of Romania as well as the soft tissue anatomy, ecology and feeding behaviour of Thalassodromeus. The specimen is actually not a pterosaur premaxillary crest at all but part of the plastron of a turtle. There is no way that 20 authors were needed to make this point but… I was one of many who collaborated on a rebuttal to the proposal.

Dyke, G. J., Vremir, M., Brusatte, S., Bever, G., Buffetaut, E., Chapman, S., Csiki-Sava, Z., Kellner, A. W. A., Martin, E., Naish, D., Norell, M., Ősi, A., Pinheiro, F. L., Prondvai, E., Rabi, M., Rodrigues, T., Steel, L., Tong, H., Vila Nova, B. C. & Witton, M. 2014. Thalassodromeus sebesensis – a new name for an old turtle. Comment on “Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur”, Grellet-Tinner and Codrea. Gondwana Research 27, 1680-1682. [pdf here]

Montage depicting the supposed premaxillary crest of ‘Thalassodromeus sebesensis‘ at left; T. sethi at lower right.

CASSOWARIES ARE AIRHEADS

As of 2014, surprisingly little had been published on cassowary evolution and anatomy, and specifically about the anatomy of the casque. Richard Perron obtained and sectioned the head of a frozen cassowary; we opted to publish a description and combine it with thoughts on cassowary phylogenetics (based on novel DNA data) and evolutionary history. Subsequent work on cassowary anatomy and physiology (including by Danielle Eastick et al., published 2019, and Todd Green and Paul Gignac, published 2020) updates and augments our conclusions.

Naish, D. & Perron, R. 2014. Structure and function of the cassowary’s casque and its implications for cassowary history, biology and evolution. Historical Biology 10.1080/08912963.2014.985669 [pdf here]

Our montage depicting casque anatomy – see Naish & Perron (2014) for explanation. Note the big air space in the casque (visible in D) and the mass of trabeculae filling its anterior part (visible in B).

• A Tetrapod Zoology write-up of this research is here: Inside the Cassowary’s Casque (February 2015)

THE SHANKLIN CROCODYLIFORM

An unusual partial dentary tip from the Upper Greensand Formation of Shanklin on the Isle of Wight (previously suggested to be from a plesiosaur) proved to be from a crocodyliform, most likely a dyrosaurid-like tethysuchian. As such, it potentially extends the distribution of these animals back in time, from the Maastrichtian to the Aptian or Albian. The specimen is anatomically unusual and likely represents a new taxon (for now, informally termed the ‘Shanklin croc’), albeit one we opted not to name at this time. Hopefully, more complete remains will one day be discovered.

Young, M. T., Steel, L., Foffa, D., Price, T., Naish, D. & Tennant, J. P. 2014. Marine tethysuchian crocodyliform from the ?Aptian-Albian (Lower Cretaceous) of the Isle of Wight, UK. Biological Journal of the Linnean Society 113, 854-871. [pdf here]

The Shanklin croc specimen (NHMUK PV OR36173) in (at top) right lateral, (at middle) ventral, and (at bottom) dorsal view. Images from Young et al. (2014). It’s not the prettiest fossil in the world, but it’s still interesting.

• A Tetrapod Zoology write-up of this research is here: ‘Shanklin croc’ and the dawn of the tethysuchian radiation

PERVASIVE SIZE DECREASE IN THEROPOD EVOLUTION

By combining a large phylogenetic analysis of Theropoda (compiled by Andrea Cau) with data on body size and geological time, we used statistical techniques to analyse size change in theropods across the course of their evolutionary history. While numerous lineages demonstrate size increase, the main pattern across the group – as determined by the body size estimated for species present at key nodes – is one of continual size decrease, from ancestral neotheropods all the way to the ancestry of birds.

Lee, M. S. Y., Cau, A., Naish, D. & Dyke, G. J. 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science 345, 562-565. [pdf here]

Theropods display a continuous, pervasive decrease in size when we look at the inferred size of ancestral species at successive nodes across the lineage leading to birds. From left to right, this illustration by Davide Bonnadonna shows the ancestral neotheropod (~220 Million years old), the ancestral tetanuran (~200 myo), the ancestral coelurosaur (~175 myo), the ancestral paravian (~165 myo), and Archaeopteryx (150 myo).

• A Tetrapod Zoology write-up of this research is here: 50 million years of incredible shrinking theropod dinosaurs (July 2014)

THE CRETACEOUS ICHTHYOSAUR REVOLUTION, PART 3

The conventional view that virtually all Cretaceous ophthalmosaurid ichthyosaurs should be absorbed into Platypterygius is not supported by phylogenetic analysis or the detailed anatomy of the specimens. In this study, led by Valentin Fischer, we examined Russian ophthalmosaurids previously lumped into Platypterygius and re-established their distinct status. We also argued that the taxa concerned have very unusual nostrils.

Fischer, V., Arkangelsky, M. S., Naish, D., Stenshin, I. M., Uspensky, G. N. & & Godefroit, P. 2014. Simbirskiasaurus and Pervushovisaurus reassessed: implications for the taxonomy and cranial osteology of Cretaceous platypterygiine ichthyosaurs. Zoological Journal of the Linnean Society 171, 822-841. [pdf here]

Holotype skull of Simbirskiasaurus birjukovi Ochev and Efimov, 1985 in left lateral view. Image from Fischer et al. (2014).

• A Tetrapod Zoology write-up of this research is here: Ancient Marine Reptiles Had Absurd, Complex Nostrils (July 2014)

A MID-CRETACEOUS ORIGIN FOR CROWN BIRDS?

Lee, M. S. Y., Cau, A., Naish, D. & Dyke, G. 2014. Morphological clocks in paleontology, and a Mid-Cretaceous origin of crown Aves. Systematic Biology 63, 442-449. [pdf here]

‘MYSTERY BIG CATS’ IN THE AMAZON

For years, reference has been made to Peter Hocking’s ‘anomalous jaguars’ from Peru, represented by skulls which Hocking obtained from local hunters. By obtaining copies of the skull and subjecting them to morphometric analysis, we were able to resolve the affinities of these previously enigmatic specimens.

Naish, D., Sakamoto, M., Hocking, P. & Sanchez, G. 2014. ‘Mystery big cats’ in the Peruvian Amazon: morphometrics solve a cryptozoological mystery. PeerJ 2:e291; DOI 10.7717/peerj.291

(A) ‘Anomalous jaguar and (B) ‘Peruvian tiger’ skulls in oblique dorsolateral view to show the flexed, concave dorsolateral part of the maxilla present in both specimens. Photo by Barry Marsh, from Naish et al. (2014).

• A Tetrapod Zoology write-up of this research is here: Mystery big cat skulls from the Peruvian Amazon not so mysterious anymore (March 2014)
• An Interview at PeerJ on this research is here

A NEW LATE CRETACEOUS VERTEBRATE ASSEMBLAGE FROM ROMANIA

Vremir, M., Bălc, R., Csiki-Sava, Z. Brusatte, S. L., Dyke, G., Naish, D., Norell, M. A. 2014. Petreşti-Arini – An important but ephemeral Upper Cretaceous continental vertebrate site in the southwestern Transylvanian Basin, Romania. Cretaceous Research 49, 13-38. [pdf here]

THE BEHAVIOUR OF FOSSIL BIRDS

Form-function correlation in extant birds is extensively studied and relatively well understood. But what do we understand about the ecology and lifestyle of fossil birds? In this review I aimed to summarise published views on current models of extinct bird lifestyle, most of which is informed by form-function correlation, though rare stomach contents and other pieces of evidence sometimes provide additional data. Some specific areas of controversy remain; also interesting is that some birds – living and fossil – appear to violate the ‘form-function’ rule.

Naish, D. 2014. The fossil record of bird behaviour. Journal of Zoology 292, 268-280. [pdf here]

Anatomical features present in fossil birds allow us to make inferences about behaviour. (a) Probable wading stem-flamingo Juncitarsus; (b) potoo-like Paraprefica; (c) marine, pseudo-toothed Pelagornis; (d) vertebrate predator Phorusrhacos; (e) slender-billed forager Rhychaeites; (f) icterid-like gaper Chascacocolius (surprisingly, a mousebird). Images not to scale. From Naish (2014).

• A Tetrapod Zoology write-up of this research is here: Bird behaviour, the ‘deep time’ perspective (January 2014)

A TINY ORNITHOPOD DINOSAUR BONE

Brusatte, S. L., Vremir, M., Watanabe, A., Csiki-Sava, Z., Naish, D., Dyke, G., Erickson, G. M. & Norell, M. A. 2013. An infant ornithopod dinosaur tibia from the Late Cretaceous of Sebeş, Romania. Terra Sebus. Acta Musei Sabesiensis 5, 627-644. [pdf here]

TESTING FLIGHT IN MICRORAPTOR

In a collaborative project involving engineers, aerodynamics experts and palaeontologists, we constructed a life-sized model of the Early Cretaceous maniraptoran theropod Microraptor (involving real feathers) and subjected it to aerodynamic tests in one of the University of Southampton wind tunnels. We aimed to test several different hindlimbs configurations for Microraptor in order to see how they affected flight performance. Our results indicate that Microraptor was not an efficient flier.

Dyke, G., de Kat, R., Palmer, C., van der Kindere, J., Naish, D. & Ganapathisubramani, B. 2013. Aerodynamic performance of the feathered dinosaur Microraptor and the evolution of feathered flight. Nature Communications 4, Article number: 2489 doi:10.1038/ncomms3489 [pdf here]

Our Microraptor – it’s called Maurice – in picturesque settings. The hindlimbs are shown here in an unrealisitic near-maximum sprawl. Photo from Dyke et al. (2013).

• A Tetrapod Zoology write-up of this research is here: Flight of the Microraptor (November 2013)

THE CRETACEOUS ICHTHYOSAUR REVOLUTION, PART 2

In the second paper resulting from the Appleby Legacy Project (see the Part 1 text on the Cretaceous Ichthyosaur Revolution below), we described and named the new Early Cretaceous ichthyosaur Malawania anachronus Fischer et al., 2013. Malawania was collected from Iraq in 1952 and was studied for years by Robert Appleby, who aimed to publish his work on it but was defeated by contradictory data on the specimen’s age. Its anatomy indicated that it was of ‘Early Jurassic grade’ yet the microfossil data he obtained indicated an Early Cretaceous age. We collected and analysed new samples and pinned down the specimen’s age to Hauterivian or Barremian (Early Cretaceous). Phylogenetic analysis showed that Malawania is part of the same lineage as Ichthyosaurus and hence demonstrates survival of its lineage (Ichthyosauridae) into the Cretaceous.

Fischer, V., Appleby, R. M., Naish, D., Liston, J., Riding, J. B., Brindley, S. & Godefroit, P. 2013. A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs. Biology Letters 9, 20130021 [pdf here]

Bob Nicholls’s life restoration of Malawania, coloured by C. M. Kosemen.

• A Tetrapod Zoology write-up of this research is here: Malawania from Iraq and the Cretaceous Ichthyosaur Revolution (part II) (May 2013)

A ‘BRITISH BIG CAT’ (A LYNX) IS SUBJECTED TO DETAILED STUDY

For years, people claim to have observed non-native cat species in the British countryside. It turns out that a lynx was living wild in Devon in the early 1900s, was shot dead, and was then accessioned in the collections of Bristol Museum. In 2010, the specimen was noticed by Max Blake, who realised its potential significance. We subjected the specimen to morphometric analysis, analysed its DNA, and used isotope analysis in an effort to determine its original provenance. This study received a substantial amount of media coverage. It is clear that more work could be done on the specimen. Its skeletal pathologies remain to be properly described, we still need to x-ray the taxiderm mount to determine its construction (we think it was done without use of the skeleton using an innovative technique), and there remains some question about the specimen’s subspecies-level identification.

Blake, M., Naish, D., Larson, G., King, C. L., Nowell, G., Sakamoto, M. & Barnett, R. 2013. Multidisciplinary investigation of a ‘British big cat’: a lynx killed in southern England c. 1903. Historical Biology doi:10.1080/08912963.2013.785541 [pdf here]

Different views of Ab4458: Canada lynx or Bobcat? Photos by Max Blake. From Blake et al. (2013).

How do the Strontium isotope values from our lynx compare to those of possible areas of residence? Annoyingly, the results are ambiguous and consistent with both western Canadian origins and with a mix of Dartmore granite/local marine sediments. From Blake et al. (2013).

Skull of Ab4458 in (A) right lateral and (B) anterior views. From Blake et al. (2013).

• A Tetrapod Zoology write-up of this research is here: A lynx, shot dead in England in c. 1903 (April 2013)
• Durham University coverage: Museum find proves exotic ‘big cat’ prowled British countryside a century ago (April 2013)

DINOSAURS AND ‘SPECIES RECOGNITION’

Palaeontologists specialising in dinosaurs have historically shown preference for the idea that extravagant structures function in ‘species recognition’. Studies on extant animals have mostly failed to show support for the species recognition hypothesis (SRH), the structures instead appearing to have evolved until sexual selection pressure. In this project – a ‘descendant’ of Knell et al. (2012) and Hone et al. (2011) – Dave Hone and I argued that the SRH should be retired as an explanation for the evolution of extravagant structure in these animals. Our study solicited a response from Kevin Padian and Jack Horner.

Hone, D. W. E. & Naish, D. 2013. The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non-avialan dinosaurs. Journal of Zoology doi:10.1111/jzo.12035 [pdf here]

A selection of flamboyant ornithischian dinosaurs. Why were so many dinosaurs so very flamboyant? So that they could tell each other apart, or because – like many living flamboyant animals – their evolution was dominated by sexual selection? Image by Darren Naish.

• A Tetrapod Zoology write-up of this research is here: Dinosaurs and their exaggerated structures: species recognition aids, or sexual display devices? (April 2013)

Knell, R. J., Naish, D., Tomkins, J. L. & Hone, D. W. E. 2013. Is sexual selection defined by dimorphism alone? A reply to Padian and Horner. Trends in Ecology and Evolution xx. [pdf here]

A NEW SMALL AZHDARCHOID PTEROSAUR, VECTIDRACO

In 2012, 4-year-old Daisy Morris found the well-preserved pelvis of a small azhdarchoid pterosaur in the Greensand rocks of the Isle of Wight. The fossil proved to represent a new taxon – Vectidraco daisymorrisae Naish et al., 2013 – and one of the world’s smallest azhdarchoids. In this initial descriptive paper, we demonstrated tapejarid affinities for Vectidraco, described its anatomy, and aimed to provide a list of pelvic character states which might be incorporated into, and coded for, in subsequent phylogenetic studies of Pterosauria. The fact that a young girl discovered an entirely new fossil vertebrate species meant that this study received an extraordinary amount of media coverage. Some anatomical errors made in this paper were corrected in Martin-Silverstone et al. (2018).

Naish, D., Simpson, M. I. & Dyke, G. J. 2013. A new small-bodied azhdarchoid pterosaur from the Lower Cretaceous of England and its implications for pterosaur anatomy, diversity and phylogeny. PLoS ONE 8(3): e58451. doi:10.1371/journal.pone.0058451

Speculative silhouette of Vectidraco as the whole animal might have looked; image by Darren Naish, from Naish et al. (2013).

• A Tetrapod Zoology write-up of this research is here: Daisy’s Isle of Wight Dragon and why China has what Europe does not (March 2013)
• PLoS BLOGS coverage: Meet Vectidraco, a European pterosaur the size of a crow (March 2013)
• National Geographic coverage: Crow-sized pterosaur named after 9-year-old fossil hunter (March 2013)

A NEW ‘MID-SIZED’ EUROPEAN AZHDARCHID PTEROSAUR

In another project resulting from our collaborative project on the vertebrate palaeontology of the Romanian Late Cretaceous, we described a new azhdarchid – Eurazhdarcho langendorfensis Vremir et al., 2013 – represented by cervical vertebrae and part of the wing skeleton. The specimen is represented by adult remains and is not a juvenile specimen of the much larger Romanian azhdarchid Hatzegopteryx. The presence of both of these azhdarchids in the same geological unit (the Sebeş Formation of the Transylvanian Basin) is significant, since it indicates that niche partitioning might have been present in sympatric azhdarchids. We noted the presence of several locations worldwide where sympatric azhdarchids occur, these taxa differing in size and hence presumably in ecology and habits. Eurazhdarcho is another new taxon discovered by Mátyás Vremir (who died in 2020).

Vremir, M., Kellner, A. W. A., Naish. D. & Dyke, G. J. 2013. A new azhdarchid pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: implications for azhdarchid diversity and distribution. PLoS ONE 8(1): e54268. doi:10.1371/journal.pone.0054268

Some geological units reveal evidence of two or even three sympatric azhdarchid species. Diagram produced by Mark Witton and map used with kind permission of Ron Blakey, Colorado Plateau Geosystems, Inc; from Vremir et al. (2013).

• A Tetrapod Zoology write-up of this research is here: A new azhdarchid pterosaur: the view from Europe becomes ever more interesting (January 2013)

ESTUARINE AND FRESHWATER PLESIOSAURS: THE LEPTOCLEIDIDS

In 1995, the partial but mostly articulated remains of a small plesiosaur were discovered in the Lower Cretaceous Vectis Formation of the Isle of Wight, UK. This animal was initially interpreted as a new specimen of the Wealden plesiosaur Leptocleidus but numerous anatomical differences suggested distinct status. In the final published paper, we determined that this animal – the ‘Shepherd’s Chine leptocleidid’ – is a new taxon (Vectocleidus pastorum Benson et al., 2012). We compared it to other leptocleidid taxa and also used the paper to analyse the position of Leptocleididae within Plesiosauria, since there are conflicting views on this issue (our data supports the proposal that leptocleidids are within Cryptoclidia and are close kin of the polycotylids, as proposed by Benson & Ketchum 2011). A second plesiosaurian taxon – Hastanectes valdensis (Benson et al. 2012) from the Hastings Group – was also named in our paper.

Benson, R. B. J., Ketchum, H. F., Naish, D. & Turner, L. E. 2012. A new leptocleidid (Sauropterygia, Plesiosauria) from the Vectis Formation (Early Barremian-early Aptian; Early Cretaceous) of the Isle of Wight and the evolution of Leptocleididae, a controversial clade. Journal of Systematic Palaeontology DOI: 10.1080/14772019.2011.634444 [pdf here]

The two more informative blocks containing the Vectocleidus pastorum holotype. We have much of the dorsal vertebral column of the animal and at least part of its pectoral skeleton. From Benson et al. (2012).

The favoured plesiosaur phylogeny from Benson et al. (2012). Note that leptocleidians are within Cryptoclidia, within Plesiosauroidea.

• A Tetrapod Zoology write-up of this research is here: Plesiosaurs and the repeated invasion of freshwater habitats: late-surviving relicts or evolutionary novelties? (January 2013)

CLAW CURVATURE AND CLIMBING IN BIRDS AND OTHER DINOSAURS

Authors from the 1980s onwards have used pedal claw shape to help determine the lifestyle and ecology of extinct birds and other animals. This technique was initially devised by Alan Feduccia to test the behaviour of Archaeopteryx (he effectively set out to prove that Archaeopteryx was arboreal or scansorial). We aimed to expand the use of this technique as much as possible in order to test its reliability with the largest dataset we could. Our results revealed a substantial amount of overlap between ecological categories and a large ‘grey area’ in between extremes.

Birn-Jeffery, A. V., Miller, C. E., Naish, D., Rayfield, E. J., Hone, D. W. E. 2012. Pedal claw curvature in birds, lizards and Mesozoic dinosaurs – complicated categories and compensating for mass-specific and phylogenetic control. PLoS ONE 7(12): e50555. doi:10.1371/journal.pone.0050555

Box plots showing inner claw curvature distinguished by behavioural category (Fig. 3 from Birn-Jeffery et al. 2012). On the left we see data from all extant taxa measured for the study, and – on the right – birds only. Note that both sets of plots are approximately similar. The most obvious thing is the massive overlap between the categories. Medians are marked by horizontal lines.

Inner claw curvature (Y axis) plotted against claw mid-length height in all extant animals (at left) and just modern birds (at right), with 22 different Mesozoic theropods added. The substantial overlap between the different behavioural categories is, again, obvious. See text (or the paper itself!) for discussion. Apologies about typo (‘Alvarezsauroidae’ should be Alvarezsauroidea).

• A Tetrapod Zoology write-up of this research is here: Did Velociraptor and Archaeopteryx climb trees? Claws and climbing in birds and other dinosaurs (December 2012)

THE ‘JAWS’ CARCASS

An analysis of a photo implied to show an unknown or new Australian mammal species (perhaps a living thylacoleonid) enabled analysis of tooth shape and distribution; sadly, the carcass is not of an unknown or new species but a familiar one. A pdf of this paper does not yet exist but I aim to create and upload one soon (August 2020).

Naish, D. 2012. Identifying ‘Jaws’, the Margaret River mammal carcase. The Journal of Cryptozoology 1, 45-55.

‘Jaws’, the Margaret River mammal carcass. See Naish (2012).

• A Tetrapod Zoology write-up of this research is here: The inaugural issue of The Journal of Cryptozoology (December 2012)

THE CADBOROSAURUS WARS, PART X

Our 2011 argument (McCormick et al. 2011) about William Hagelund’s ‘baby cadborosaurus’ – a key piece of evidence in Bousfield and LeBlond’s argument about what Cadborosaurus is like and what sort of animal it is – was challenged by Bousfield and LeBlond in a brief 2012 paper. They argued that we were guilty of “home-quarterbacking” (I think this is a sportsing reference) and that our pipefish proposal was less likely than their alternative: that Hagelund’s creature was indeed a baby cadborosaur. We thought that their phrasing of our argument was unfair and reiterated our case in this response.

Woodley, M. A., McCormick, C. A. & Naish, D. 2012. Response to Bousfield & LeBlond: Shooting pipefish in a barrel; or sauropterygian “mega-serpents” and Occam’s razor. Journal of Scientific Exploration 26, 151-154.

• A Tetrapod Zoology write-up of this research is here: The Cadborosaurus Wars (April 2012)

SEXUAL SELECTION AND FOSSILS, OPENING SALVO

That sexual selection is one of the primary drivers in evolution is well established. Studies of Mesozoic archosaurs (dinosaurs and pterosaurs), however, have been dominated by the view that the variation, extravagance and diversity seen in these animals is not due to the pressures of sexual selection, but to other evolutionary phenomena. In this 2012 article we sought to argue that there are good grounds for promoting sexual selection as a driver in the evolution of extravagant structures in fossil animals in general. This article can be seen as the ancestor of several subsequent, related projects. TREE is an extremely influential journal, with an Impact Factor of 14.7!

Knell, R. J., Naish, D., Tomkins, J. L. & Hone, D. W. E. 2012. Sexual selection in prehistoric animals: detection and implications. Trends in Ecology and Evolution 28, 38-47. [pdf here]

Select fossil vertebrates with ‘exaggerated morphologies’, from Knell et al. (2012). a, Achelousaurus. b, Olorotitan. c, Protoceras. d, Akmonistion. e, Diceratosaurus. f, Diplocaulus. Images by Darren Naish (redrawn from various sources. Not to scale).

• A Tetrapod Zoology write-up of this research is here: Sexual Selection in the Fossil Record (September 2012)
• Survival of the prettiest: sexual selection can be inferred from the fossil record (ScienceDaily, January 2013)

AN ENANTIORNITHINE NESTING COLONY, PART 1

As part of our collaborative project on the vertebrate palaeontology of the Romanian Late Cretaceous, myself and colleagues described an eggshell-dominated sedimentary lens discovered by Mátyás Vremir. The eggshell was mostly that produced by enantiornithines, so here is evidence that some enantiornithines nested in large waterside colonies much like those of modern gulls, plovers and waders. An update to this research was published in Fernández et al. (2019)

Dyke, G., Vremir, M., Kaiser, G. & Naish, D. 2012. A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania. Naturwissenschaften 99, 435-442. [pdf here]

• A Tetrapod Zoology write-up of this research is here: A drowned nesting colony of Late Cretaceous birds (May 2012)

THE CRETACEOUS ICHTHYOSAUR REVOLUTION, PART 1

A collaborative effort led by Jeff Liston and the late Arthur Cruickshank (1932-2011) has revolved around efforts to salvage for publication the work of former ichthyosaur worker Robert Appleby (1924-2004); it is known as the Appleby Legacy Project. Several of the ichthyosaurs under study by Appleby have special relevance to our understanding of ichthyosaur diversity and evolution in the Cretaceous; we informally speak of the relevant studies driving an event we termed the Cretaceous Ichthyosaur Revolution. In this, the first of our several relevant papers, we described the new Early Cretaceous ophthalmosaurid Acamptonectes densus Fischer et al., 2012. This taxon is significant in demonstrating the persistence of the Ophthalmosaurus lineage into the Cretaceous.

Fischer, V., Maisch, M. W., Naish, D., Kosma, R., Liston, J., Joger, U., Krüger, F. J., Pérez, J. P., Tainsh, J. & Appleby, R. M. 2012. New ophthalmosaurid ichthyosaurs from the European Lower Cretaceous demonstrate extensive ichthyosaur survival across the Jurassic-Cretaceous boundary. PLoS ONE 7(1): e29234. doi:10.1371/journal.pone.0029234

The new Hauterivian ophthalmosaurine ichthyosaur Acamptonectes densus Fischer et al., 2012, as reconstructed by C. M. Kosemen.

• A Tetrapod Zoology write-up of this research is here: Rigid Swimmer and the Cretaceous Ichthyosaur Revolution (part I) (January 2012)
• An article by Jeff Liston on our Acamptonectes paper: Missing Skull-Bones: Hidden Sea Dragon – A guest post by Jeff Liston (January 2012)

Dyke, G. J. & Naish, D. 2011. What about European alvarezsauroids Proceedings of the National Academy of Sciences of the United States of America doi:10.1073/pnas.1101602108 [pdf here]

A NEW TINY MANIRAPTORAN FROM THE WEALDEN

In another project on the Early Cretaceous Wealden theropods of southern England, Steve Sweetman and I described a tiny cervical vertebra from a theropod which Steve discovered in the Hastings Group of Ashdown, East Sussex. The vertebra’s total length is just 7.1 mm. It appears to be from an oviraptorosaur-type animal; estimates suggest a total length for this animal of perhaps 33 cm. This makes it among the smallest non-bird theropods yet reported. It also highlights the fact that much remains unknown about small theropod diversity in the British Early Cretaceous. This paper received an absurd amount of media coverage, even though we didn’t put out at a press release (my blog article was sufficient to drum up interest).

Naish, D. & Sweetman, S. C. 2011. A tiny maniraptoran dinosaur in the Lower Cretaceous Hastings Group: evidence from a new vertebrate-bearing locality in south-east England. Cretaceous Research 32, 464-471. [pdf here]

• A Tetrapod Zoology write-up of this research is here: A truly tiny Cretaceous theropod… from England? (May 2011)

SAUROPODS NECKS WERE ‘NOT FOR SEX’

A 1996 claim that giraffe neck evolved under sexual selection pressure led Phil Senter to argue that the remarkable necks of sauropod dinosaurs might also have evolved as sexual display structures. Senter’s 2007 study, published in Journal of Zoology, became one of the journal’s most-downloaded papers. Myself and colleagues saw several issues with Senter’s proposal and sought to contest it in a 2011 response. Phil Senter emailed us afterwards to say that he agreed with our interpretation of the data, and thanked us for a fair and thorough evaluation of his proposal. It should be noted that our argument does not negate the potential use of the sauropod neck in sexual display; rather, we argue that sexual selection was not the primary driver of neck size in these animals.

Taylor, M. P., Hone, D. W. E., Wedel, M. J., & Naish, D. 2011. The long necks of sauropods did not evolve primarily through sexual selection. Journal of Zoology 285, 150-161. [pdf here]

Sauropods (like these Sauroposeidon) might well have used the neck in sexual display, but it doesn’t follow that sexual selection explains the evolution of their necks. This image is (c) Brian Engh and comes from dontmesswithdinosaurs.com.

• A Tetrapod Zoology write-up of this research is here: Necks for sex? No thank you, we’re sauropod dinosaurs (May 2011)
• SV-POW!: Why the long necks? Probably not sexual selection (May 2011)
• Riley Black at SmithsonianMag.com covered this research: Sex and Dinosaur Necks (February 2012)

THE CADBOROSAURUS WARS, PART X

Woodley, M. A., Naish, D., & McCormick, C. A. 2011. A baby sea-serpent no more: reinterpreting Hagelund’s juvenile “cadborosaur” report. Journal of Scientific Exploration 25, 495-512. [pdf here]

DINOSAURS, PTEROSAURS AND THE MUTUAL SEXUAL SELECTION

Non-bird dinosaurs and pterosaurs are well known for the diversity of their extravagant structures, which include crests, casques, horns, dorsal sails, spikes and spines. If these structures are interpreted as sexual display structures, it might follow that they display sexual dimorphism. Several authors have claimed to find dimorphism in these animals (variously in hadrosaurs, ceratopsians and stegosaurs) but all cases are problematic, mostly because supposed male and female data-points overlap and appear to exhibit continuous variation. A possibility which hadn’t been considered by previous authors is that these animals might have exhibited mutual sexual selection, a phenomenon in which males and female are similarly extravagant.

Hone, D. W. E., Naish, D. & Cuthill, I. C. 2011. Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs? Lethaia 45, 139-156. [pdf here]

A selected of crested pterodactyloid pterosaurs. A. Ornithocheirid ornithocheiroid Ornithocheirus. B. Tapejarid azhdarchoid Tapejara. C. Thalassodromid azhdarchoid Tupuxuara. D. Dsungaripterid Dsungaripterus. E. Pteranodontid ornithocheiroid Pteranodon. F. Nyctosaurid ornithocheiroid Nyctosaurus. Diagrams by Dave Hone.

• A Tetrapod Zoology write-up of this research is here: Did dinosaurs and pterosaurs practise mutual sexual selection?  (January 2012)

SAMRUKIA: BECAUSE WE ALL MAKE MISTAKES

In August 2010, I was given access to a seemingly remarkable large lower jaw from a Late Cretaceous animal, the toothless, U-shaped symphysis of which made it look like an oviraptorosaurian or bird mandible. Even after the symphysis was removed (it proved to be reconstructed in plaster), I persisted with the theropod identification (specifically identifying it as a bird), and this was supported by phylogenetic analysis and via the opinions of colleagues. We thus published this specimen as a new taxon – Samrukia nessovi Naish et al., 2012 – of surprisingly large archaic bird. I had overlooked the fact that the articular region was clearly that of a pterosaur, and a paper reidentifying Samrukia as a pterosaur was published almost immediately by Eric Buffetaut. However, this is not the end of the story and I aim to publish a follow-up paper when time allows.

Naish, D., Dyke, G., Cau, A., Escuillié, F. & Godefroit, P. 2012. A gigantic bird from the Upper Cretaceous of Central Asia. Biology Letters 8, 97-100. [pdf here]

Two possible body shapes for the gigantic Samrukia (back when we incorrectly interpreted it as a bird), with a human and ‘normal-sized’ Mesozoic bird for scale. Image by John Conway.

• A Tetrapod Zoology write-up of this research is here: Big birds in the Cretaceous of Central Asia: say hello to Samrukia (August 2011)

THE CORNET BIRDS AND PTEROSAURS

A number of small pterosaur and bird bones from the Lower Cretaceous of Cornet in Romania – currently housed at the Tarii Crisurilor Museum, Oradea – have been given a number of very specific identifications over the years: some were identified as elements of archaeopterygids or crown-birds. We evaluated these proposals and suggested new identifications.

Dyke, G., Benton, M., Posmosanu, E., & Naish, D. 2010. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology 54, 79-95. [pdf here]

• A write-up of this paper was covered in two Tetrapod Zoology articles. Part I is here, Part II is here.

CAN GIRAFFES SWIM?

Several books and article states that giraffes cannot swim. I became interested in testing this contention. I knew that Don Henderson had produced a digital model of a giraffes and was able to test the buoyancy and swimming abilities of his models via digital simulation (he had done it previously for various dinosaurs). Would it be possible for him to ‘float’ his digital giraffe and test the swimming dynamics of giraffes once and for all? This paper was the result of our experiment. The project received a substantial amount of media coverage, was the focus of a brief Scientific American article, and was even discussed on QI in 2020.

Henderson, D. M. & Naish, D. 2010. Predicting the buoyancy, equilibrium and potential swimming ability of giraffes by computational analysis. Journal of Theoretical Biology 265, 151-159. [pdf here]

• A Tetrapod Zoology write-up of this research is here: Testing the flotation dynamics and swimming abilities of giraffes by way of computational analysis (June 2010)
• BBC Earth article which discusses this research: The strange experiments that revealed that most mammals can swim (March 2017)
• Treehugger article: Think Giraffes Can’t Swim? Science Proves They Can (October 2018)

SAUROPOD NECKS WERE LIKELY HELD ERECT

In 1999, Kent Stevens and Mike Parrish published an influential Science paper in which they argued that the long necks of diplodocid sauropods were likely held in near-horizontal poses, and that the nature of bone-bone articulations in the vertebrae of these dinosaurs showed that erect or semi-erect neck poses were not likely. This explains why the diplodocids in the 1999 BBC TV series Walking With Dinosaurs have straight, horizontal necks. In a series of subsequent papers, Stevens and Parrish extended their model to Euhelopus, Brachiosaurus, Camarasaurus, Dicraeosaurus… effectively to all sauropods. The problem with this research is that it assumes that direct bone-to-bone articulation provides a reliable guide to life posture of the neck, and this is inconsistent with data from live animals (which of course can be observed, alive, in x-ray). We used data from a diversity of live tetrapods to show that the assumed resting pose of an animal – even one with a ridiculously long neck, like a sauropod – should be assumed to involve strong extension at the neck base, strong flexion at the head-neck junction, and a mostly straight ‘central’ portion. This paper generated a huge amount of media coverage and much discussion in the dinosaur research community. Kent Stevens published a response paper and several conference talks have been devoted to this area of disagreement (which has informally been termed the ‘neck wars’).

Taylor, M. P., Wedel, M. J. & Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54, 213-220. [pdf here]

• A Tetrapod Zoology write-up of this research is here: Sauropod dinosaurs held their necks in high, raised postures (May 2009)
• ScienceDaily: Giant Dinosaur Posture Is All Wrong: Sauropods Held Their Heads High, Research Finds (May 2009)
• Riley Black at SmithsonianMag.com: The Sauropod Posture Debate, Part Eleventy (May 2009)
• MailOnline: Why dinosaurs can hold their heads high after scientists say classic portrayal is wrong (May 2009)
• SV-POW! list of media coverage: Taylor, Wedel and Naish (2009) on neck posture
• Mike Taylor’s site: Sauropods held their necks erect (March 2011)

A NEW LARGE THEROPOD DINOSAUR FROM THE WEALDEN OF THE ISLE OF WIGHT

During the course of my PhD work at the University of Portsmouth I encountered the remains of a new Wessex Formation theropod, which I wrote up (in my thesis) as a new taxon of uncertain affinities. It turned out that other bones belonging to the same specimen were known and being worked on by a second group of authors. Fortunately this was realised before either of us went to print. The result was our brief description of the ‘6350 theropod’, a mid-sized taxon, distinct from named Wessex Formation theropods.

Benson, R. B. J., Brusatte, S. L., Hutt, S. & Naish, D. 2009. A new large basal tetanuran (Dinosauria: Theropoda) from the Wessex Formation (Barremian) of the Isle of Wight, England. Journal of Vertebrate Paleontology 29, 612-615. [pdf here]

• A Tetrapod Zoology write-up of this research is here: Oh no, not another new Wealden theropod! (June 2009)

AZHDARCHIDS WERE TERRESTRIAL STALKERS

Azhdarchids have to be regarded as the most distinctive and unusual of pterosaur groups, and views on how they might have lived have varied considerably. Mark Witton and I examined the skeletal proportions, cranial anatomy and sedimentological setting of azhdarchids and concluded that they were strongly adapted for quadrupedal walking in terrestrial settings. They were likely striding predators of small and mid-sized prey, analogous to modern ground hornbills. We termed this the ‘terrestrial stalking’ model. Our proposal has been supported by additional studies (Witton & Naish 2015, Naish & Witton 2017) though there has been some pushback from other pterosaur specialists.

Witton, M. P. & Naish, D. 2008. A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS ONE 3 (5): e2271. doi:10.1371/journal.pone.0002271

Image (c) Mark Witton.

• A Tetrapod Zoology write-up of this research is here: Terrestrial stalking azhdarchids, the paper (May 2008)

ARE THERE MORE PINNIPEDS TO DISCOVER?

In this study – inspired by work which uses discovery rates to estimate the number of big extant vertebrates possibly awaiting discovery – we looked at pinniped discovery rates over time. Data supports the possibility that a very low number might still be out there, but the number is so low that it could also be 0! We also evaluated some supposed crypto-pinnipeds of the cryptozoological literature. This study has been cited in the cryptozoological literature as support for the idea that new pinnipeds might be out there to find, and as support for the idea that there are no new pinnipeds out there to find.

Woodley, M. A., Naish, D. & Shanahan, H. P. 2008. How many extant pinniped species remain to be described? Historical Biology 20, 225-235. [pdf here]

Image (c) C. M. Kosemen

• A Tetrapod Zoology write-up of this research is here: Statistics, seals and sea monsters in the technical literature (March 2009)

Naish, D. & Martill, D. M. 2008. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: Ornithischia. Journal of the Geological Society, London 165, 613-623. [pdf here]

Sánchez-Hernández, B., Benton, M. J. & Naish, D. 2007. Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 249, 180-215. [pdf here]

THE NEW WEALDEN SAUROPOD XENOPOSEIDON

As part of a long-running study on Wealden sauropods, Mike P. Taylor and I reported the new sauropod Xenoposeidon proneneukos Taylor & Naish, 2007, a Wealden sauropod represented by an odd, anatomically novel vertebra. This specimen is from the Hastings Group, the ‘old’ part of the Wealden which crops out on the English mainland (especially in East Sussex) rather than on the Isle of Wight. The single vertebra is so odd that we were initially unable to allocate it to any recognised sauropod group, and our paper implied that it might represent a new ‘family’. Later publications have shown that Xenoposeidon is actually a rebbachisaurid diplodocoid. The initial paper received a vast quantity of media coverage.

Taylor, M. P. & Naish, D. 2007. An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50, 1547-1564. [pdf here]

• A Tetrapod Zoology write-up of this research is here: The world’s most amazing sauropod (November 2007)
• A second TetZoo article, marking the 10th anniversary of this paper, is here: 10 Long, Happy Years of Xenoposeidon (November 2017)

Naish, D. & Martill, D. M. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164, 493-510. [pdf here]

Martill, D. M., Earland, S. & Naish, D. 2006. Dinosaurs in marine strata: evidence from the British Jurassic, including a review of the allochthonous vertebrate assemblage from the marine Kimmeridge Clay Formation (Upper Jurassic) of Great Britain. In Colectivo Arqueológico-Paleontológico Salense (ed) Actas de las III Jornadas sobre Dinosaurios y su Entorno. Salas de los Infantes (Burgos, España), pp. 47-83. [pdf here]

NEW DATA ON THE AZHDARCHOID PTEROSAUR TUPUXUARA

A new partial skull from the ‘mid Cretaceous’ of Brazil appears to represent the posterodorsal section of the cranial crest of Tupuxuara. Because our specimen was clearly much smaller than the holotype (and some referred specimens), we argue that it represented a younger ontogenetic stage and thus provided new information on growth changes which happened in the skulls of these pterosaurs. We used this study to evaluate the taxonomy of Tupuxuara and its relatives and argued here that the giant Thalassodromeus should be regarded as an old adult Tupuxuara, a view we later abandoned. We also used this study to evaluate the phylogeny of these pterosaurs, concluding that thalassodromids are closer to azhdarchids than to tapejarids. Pterosaur experts continue to argue on the relationship between these lineages. The publicity push for this paper involved one of the first public outings of Mark Witton artwork.

Martill, D. M. & Naish, D. 2006. Cranial crest development in the azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly. Palaeontology 49, 925-941. [pdf here]

Image (c) Mark Witton.

• A Tetrapod Zoology write-up of this research is here: Those sexy tupuxuarids (November 2006)

Fielding, S., Martill, D. M. & Naish, D. 2005. Solnhofen-style soft-tissue preservation in a new species of turtle from the Crato Formation (Early Cretaceous, Aptian) of north-east Brazil. Palaeontology 48, 1301-1310. [pdf here]

Taylor, M. P. & Naish, D. 2005. The phylogenetic taxonomy of Diplodocoidea (Dinosauria: Sauropoda). PaleoBios 25, 1-7. [pdf here]

Naish, D., Martill, D. M. & Frey, E. 2004. Ecology, systematics and biogeographical relationships of dinosaurs, including a new theropod, from the Santana Formation (?Albian, Early Cretaceous) of Brazil. Historical Biology 16, 57-70. [pdf here]

A SAUROPOSEIDON-LIKE WEALDEN SAUROPOD

During the 2000/2001 preparation of our book Dinosaurs of the Isle of Wight (Martill & Naish 2001), Dave Martill and I became aware of a surprisingly large cervical vertebra from the Wessex Formation of the Isle of Wight. It was clearly from a giant macronarian sauropod and was irresistibly similar to the cervical vertebrae of Brachiosaurus, Giraffatitan and (the then brand-new) Sauroposeidon (all of which were regarded as brachiosaurids at the time: Sauroposeidon is now regarded as belonging elsewhere within Somphospondyli). We interpreted the new specimen (a second, less well preserved vertebra is also known) as belonging to a Sauroposeidon-like animal, likely over 20 m long. As ‘Europe’s largest dinosaur’, it understandably received a massive amount of media coverage… and was toppled in 2006 by the Spanish Turiasaurus. The Wessex Formation animal likely represents a new taxon (informally dubbed ‘Angloposeidon’); additional studies on its internal structure are planned but (as of September 2020) not yet complete.

Naish, D., Martill, D. M., Cooper, D. & Stevens, K. A. 2004. Europe’s largest dinosaur? A giant brachiosaurid cervical vertebra from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 25, 787-795. [pdf here]

• Four Tetrapod Zoology articles cover this research: ‘Angloposeidon’, the unreported story, part I (July 2006); ‘Angloposeidon’, the unreported story, part II (July 2006); ‘Angloposeidon’, the unreported story, part III (July 2006); ‘Angloposeidon’, the unreported story, part IV (July 2006)

Naish, D. 2004. New Zealand’s giant gecko: a review of current knowledge of Hoplodactylus delcourti and the kawekaweau of legend. The Cryptozoology Review 4 (2), 17-21.

Naish, D. & Dyke, G. J. 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2004, 385-401. [pdf here]

Naish, D. & Martill, D. M. 2003. Pterosaurs – a successful invasion of prehistoric skies. Biologist 50 (5), 213-216. [pdf here]

ANOTHER ALLOSAUROID IN THE WEALDEN (WEALDEN THEROPODS 6)

The proximal end of a large theropod tibia was recovered from an unknown formation in the Lower Cretaceous Hastings Group. It most likely belongs to an member of the Allosauridae + Carcharodontosauria clade within Allosauroidea rather than to a metriacanthosaurid, but beyond that its affinities are unknown. The possibility that it might be referable to the (probably carcharodontosaurian) Altispinax exists, but cannot be tested without associated remains.

Naish, D. 2003. A definitive allosauroid (Dinosauria; Theropoda) from the Lower Cretaceous of East Sussex. Proceedings of the Geologists’ Association 114, 319-326. [pdf here]

THE SMALL WEALDEN THEROPODS CALAMOSPONDYLUS AND ARISTOSUCHUS (WEALDEN THEROPODS 5)

Archival research revealed that two Wealden theropods generally considered synonymous by previous researchers – Calamospondylus oweni and Aristosuchus pusillus – were based on different specimens, one of which is currently lost. This paper is mostly on historical taxonomy but I managed to include a bit of descriptive osteology in there too. The relevant articles (from the 1860s) also included reference to tree-climbing in these dinosaurs, which explains my interest in that subject, explored in other articles…

Naish, D. 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria) Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings of the Geologists’ Association 113, 153-163. [pdf here]

Speculative reconstruction of a deceased Aristosuchus, showing the relative position of its preserved elements. Image: Darren Naish.

Walsh, S. A. & Naish, D. 2002. Fossil seals from late Neogene deposits in South America: a new pinniped (Carnivora, Mammalia) assemblage from Chile. Palaeontology 45, 821-842. [pdf here]

IS THECOCOELURUS AN OVIRAPTOROSAUR? (WEALDEN THEROPODS 4)

In the fourth of my contributions on Wealden theropods, I redescribed and reintepreted Thecocoelurus from the Isle of Wight, named for just half a cervical vertebra. It looks quite similar to the cervical vertebrae of caenagnathid oviraptorosaurs, so that’s what Dave Martill and I suggested it is (a proposal backed up by phylogenetic analysis). I revisited this idea in later years, later suggested that it might be from a noasaurid.

Naish, D. & Martill, D. M. 2002. A reappraisal of Thecocoelurus daviesi (Dinosauria: Theropoda) from the Early Cretaceous of the Isle of Wight. Proceedings of the Geologists’ Association 113, 23-30. [pdf here]

EOTYRANNUS IS DESCRIBED (WEALDEN THEROPODS 3)

In this third contribution on Wealden theropods, I and colleagues preliminarily described the new Wessex Formation tyrannosauroid Eotyrannus lengi Hutt et al., 2001 from the Isle of Wight. The full monograph on this taxon would form the main substance of my PhD thesis. At the time of writing (August 2020), a full Open Access paper has been written — and really needs finishing and publishing.

Hutt, S., Naish, D., Martill, D. M., Barker, M. J. & Newbery, P. 2001. A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 22, 227-242. [pdf here]

Eotyrannus vs Hypsilophodon, as visualised by Luis Rey in 2001, (c) Luis Rey.

MORE WEALDEN THEROPOD LEG BONES (WEALDEN THEROPODS 2)

Naish, D. 2000. A small, unusual theropod (Dinosauria) femur from the Wealden Group (Lower Cretaceous) of the Isle of Wight, England. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2000, 217-234. [pdf here]

Naish, D. 2000. Theropod dinosaurs in the trees: a historical review of arboreal habits amongst nonavian theropods. Archaeopteryx 18, 35-41. [pdf here]

A WEALDEN THEROPOD TIBIA, DISCOVERED BY GIDEON MANTELL (WEALDEN THEROPODS 1)

My first peer-reviewed paper discussed a partial theropod tibia from the Wealden of East Sussex, seemingly from a mid-sized tetanuran. Associated correspondence revealed that it had been obtained by Gideon Mantell, though he seems to have misidentified it as the limb bone of a turtle. It is also marked with what look like bite marks from a theropod.

Naish, D. 1999. Theropod dinosaur diversity and palaeobiology in the Wealden Group (Early Cretaceous) of England: evidence from a previously undescribed tibia. Geologie en Mijnbouw 78, 367-373. [pdf here]

THE CAMP FIRCOM CADDY CARCASS

Naish, D. 1997. Another Caddy carcass? The Cryptozoology Review 2 (1), 26-29. [pdf here]

Naish, D. 1996. Analysing video footage purporting to show the “migo” – a lake monster from Lake Dakataua, New Britain. The Cryptozoology Review 1 (2), 18-21. [pdf here]

Book Chapters

Naish, D. 2013. South American native mammals. In McDade, M. (ed) Grzimek’s Animal Life Encyclopedia: Extinct Life. Gale Group (Farmington Mills, Michigan), pp. 567-576.

Naish, D. 2012. Birds. In Brett-Surman, M. K., Holtz, T. R. & Farlow, J. O. (eds) The Complete Dinosaur (Second Edition). Indiana University Press (Bloomington & Indianapolis), pp. 379-423.

Naish, D. 2011. Theropod dinosaurs. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 526-559.

Salisbury, S. W. & Naish, D. 2011. Crocodilians. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 305-369. [pdf here]

Naish, D. 2010. Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurischian pneumaticity. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 229-236. [pdf here]

Moody, R. T. J. & Naish, D. 2010. Alan Jack Charig (1927-1997): an overview of his academic accomplishments and role in the world of fossil reptile research. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 89-109. [pdf here]

Naish, D., Martill, D. M. & Merrick, I. 2007. Birds of the Crato Formation. In Martill, D. M., Bechly, G. & Loveridge, R. F. (eds) The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press (Cambridge), pp. 525-533.

Naish, D. 2007. Turtles of the Crato Formation. In Martill, D. M., Bechly, G. & Loveridge, R. F. (eds) The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press (Cambridge), pp. 452-457.

Books

Conway, J., Kosemen, C. M. & Naish, D. 2013. Cryptozoologicon Volume I. Irregular Books.

Conway, J., Kosemen, C. M. & Naish, D. 2012. All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals. Irregular Books.

Naish, D. 2011. Dinosaur Record Breakers. Carlton Books, London.

Naish, D. 2010. Tetrapod Zoology Book One. CFZ Press, Bideford.

Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. 2010. Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications.

Naish, D. 2010. Dinosaurs Life Size. Barron’s Educational Series, New York.

Naish, D. 2009. The Great Dinosaur Discoveries. A & C Black, London.

Martill, D. M. & Naish, D. 2001. Dinosaurs of the Isle of Wight. The Palaeontological Association, London.

Martill, D. M. & Naish, D. 2000. Walking With Dinosaurs: The Evidence. BBC Worldwide, London.

Book Reviews

Naish, D. 2015. [Review of]  The ecology and conservation of Asian hornbills: farmers of the forest. Historical Biology: An International Journal of Paleobiology 27, 954-956. [pdf here]

Naish, D. 2014. [Review of] Pterosaurs, by Mark P. Witton. Princeton University Press, Princeton and Oxford, 2013, 291 pp., ISBN 978-0-691-15061-1. Historical Biology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2014.882099 [pdf here]

Naish, D. 2014. A Review of ‘The Dodo and the Solitaire: A Natural History’. Journal of Vertebrate Paleontology 34, 489-490. [pdf here]

Naish, D. 2012. [Review of] New perspectives on horned dinosaurs: the Royal Tyrrell Museum Ceratopsian Symposium. Historical Biology doi: 10.1080/08912963.2012.688589 [pdf here]

Naish, D. 2012. [Review of] Biology of the Sauropod Dinosaurs: Understanding the Life of Giants edited by Nicole Klein, Kristian Remes, Carole T. Gee, and P. Martin Sander. The Quarterly Review of Biology 87, 53.

Naish, D. 2012. [Review of] Dinosaurs: a field guide/The Princeton field guide to dinosaurs. Historical Biology DOI:10.1080/08912963.2012.670534 [pdf here]

Naish, D. 2011. [Review of] Barnum Brown: the man who discovered Tyrannosaurus rex. Historical Biology DOI:10.1080/08912963.2011.630260 [pdf here]

Naish, D. 2011. [Review of] The second Jurassic dinosaur rush: museums and paleontology in America at the turn of the twentieth century. Historical Biology  DOI:10.1080/08912963.2011.614404 [pdf here]

Naish, D. 2011. [Review of] Glorified dinosaurs: the origin and evolution of birds. Historical Biology 23, 435-438.

Naish, D. 2011. [Review of] The inner bird: anatomy and evolution. Historical Biology 23, 313-316. [pdf here]

Naish, D. 2008. [Review of] Patagonian Mesozoic Reptiles. The Palaeontology Newsletter 62, 92-97.

Naish, D. 2006. The Carnivorous Dinosaurs [review]. The Palaeontology Newsletter 62, 122-126.

Naish, D. 2004. [Review of] African Dinosaurs Unearthed: the Tendaguru Expeditions. Proceedings of the Geologists’ Association 115, 379-380.

Naish, D. 2003. [Review of] The Interrelationships and Evolution of Basal Theropod Dinosaurs. Geological Magazine 140, 729.

Popular and Semi-Popular Articles

Naish, D. 2014. Speculative zoology. Fortean Times 316, 52-53. [pdf here]

Naish, D. 2013. How dinosaurs conquered the world. BBC Focus (Feb’ 2013) 251, 54-59.

Naish, D. 2012. Questions at the frontiers of… dinosaur science. BBC Focus October 2012, 32-33.

Naish, D. 2012. Palaeontology bites back… (response to Ford’s article on alleged aquatic habits of dinosaurs) Laboratory News May 2012, 31-32. [pdf here]

Naish, D. 2012. Should we give up looking for Bigfoot? BBC Focus March 2012, 27. [pdf here]

Naish, D. 2010. Will it float? Scientific American 304 (1), 22. [pdf here]

Naish, D. 2008. Intelligent dinosaurs. Fortean Times 239, 52-53. [pdf here]

Naish, D. 2005. Fossils explained 51: sloths. Geology Today 21 (6), 232-238. [pdf here]

Naish, D. 2004. Fossils explained 48. Placodonts. Geology Today 20 (4), 153-158. [pdf here]

Naish, D. 2004. Fossils explained 46. Ancient toothed whales. Geology Today 20 (2), 72-77. [pdf here]

Naish, D. 2002. Thecocoelurians, calamosaurs and Europe’s largest sauropod: the latest on the Isle of Wight’s dinosaurs. Dino Press 7, 85-95.

Naish, D., Noe, L. F. & Martill, D. M. 2001. Giant pliosaurs and the mysterious ‘Megapleurodon’. Dino Press 4, 98-103.

Naish, D. 2001. Eotyrannus lengi, a new coelurosaur from the Isle of Wight. Dino Press 5, 82-91.

Naish, D. 2001. Fossils explained 34: Crocodilians. Geology Today 17 (2), 71-77. [pdf here]

Naish, D. 2000. 130 years of tree-climbing dinosaurs: Archaeopteryx, ‘arbrosaurs’ and the origin of avian flight. The Quarterly Journal of the Dinosaur Society 4 (1), 20-23. [pdf here]

The tree-climbing Mesozoic theropods hypothesised to exist by certain authors: Rozhdestvensky’s sloth-like Deinocheirus, Chatterjee’s tree-climbing compsognathid (!) and ornithomimid (!!), Palm’s scansorial dromaeosaurids.

Naish, D. 2000. Where be monsters? (sea serpents). Fortean Times 132, 40-44. [pdf here]

Naish, D. 1999. Fossil Explained 27: Theropod dinosaurs. Geology Today 15, 234-239. [pdf here]

Naish, D. 1998. Birds of a feather. Fortean Times 108, 34-37.

Other Stuff

Kosemen, C. M. 2017. All Your Yesterdays (second edition). Irregular Books. [pdf here]

Simpson, M. I. 2015. Iguanodon is older than you think: the public and private announcements of Gideon Mantell’s giant prehistoric herbivorous reptile. Deposits [pdf here]

Simpson, M. I. 2020. Walk that changed history: new evidence about the discovery of the Iguanodon. Deposits [pdf here]

A Vast Quantity of Evidence Confirms That Non-Bird Dinosaurs Were Not Aquatic (supplementary info for the Brian Ford v Darren Naish debate of 15th May 2018) [pdf here]

Ings, S. 2020. What lies beneath [review of Monsters of the Deep exhibition at National Maritime Museum, Falmouth]. Financial Times Weekend Magazine 861 (March 14/15 2020), 42-47. [pdf here]

Table of new taxa co-named by Darren Naish [pdf here]