Professor Jenny Clack, 1947-2020

Within recent decades, four specific areas of palaeontological discovery and reinterpretation have succeeding in capturing mainstream attention: the feathering of dinosaurs, the evolution of hominins, the early history of whales, and the early evolution, anatomy and biology of the earliest tetrapods. And it’s the last of those subjects we’re focusing on here. It’s a subject which has seen regular airing in top-tier journals, science magazines and TV documentaries, and one which has undergone a major and exciting revolution since the 1980s.

Caption: at left, Professor Jenny Clack, in the field at Burnmouth in the Scottish Borders. At right: in 2017, Jenny and her husband Rob got to fly in a Spitfire. Images: (c) Rob Clack, used with permission.

One person above all others has been responsible for leading research in this field and has made numerous ground-breaking discoveries herself, both in the field and laboratory. I am of course referring to Professor Jenny Clack of the Department of Zoology at the University of Cambridge, an excellent and highly respected scientist whose technical papers are authoritative, ground-breaking and of the highest standard. Her publication list is formidable. Alas, I regret to write that I’m here for sad reasons, since Jenny died on the morning of 26th March 2020 following a five-year struggle with cancer. While – for my shame – I haven’t written about early tetrapod evolution here at TetZoo before, nor about Jenny’s research specifically, both are areas I’ve avidly followed, and I want to share some brief thoughts here.

Caption: a selection of Clack publications in the TetZoo library. Image: Darren Naish.

Prior to Jenny’s work, the consensus view on the oldest tetrapods – in particular Ichthyostega from the Devonian of Greenland – is that these were ‘terrestrialised’, vaguely salamander-like quadrupeds, well able to walk on land by planting all four feet on the substrate. This view, entrenched in textbooks and the popular literature, mostly owed itself to the work of Swedish palaeontologist Erik Jarvik whose work on Ichthyostega, while initiated in the 1950s, took some decades to appear, finally being published in 1996. Jarvik’s work has its own long and curious backstory (Ries 2007) – which I can’t cover here – and it certainly wasn’t missed by some of us that the conclusions of his supposedly definitive monograph were being called into doubt just weeks after its appearance (Norman 1996).  

Caption: Jenny Clack (and Professor Robert Insall) at the Ballagan Formation type locality, near Glasgow. Image: (c) Rob Clack, used with permission.

Tantalising remains of an Ichthyostega contemporary – the smaller Acanthostega – were discovered in East Greenland on a Cambridge University expedition of 1970 (and were languishing, unrecognised, in the drawers of the university’s earth sciences department prior to Jenny’s interest). These indicated that, with luck, more early tetrapod finds might be discovered in the same region. After making special arrangements with Danish scientists – and with Jarvik, who the Danes wanted to remain on good terms with – Jenny visited Greenland in 1987; she was accompanied by colleagues from Copenhagen, her husband Rob and her then PhD student Per Ahlberg (Clack 1988). They were remarkably successful, retrieving multiple good specimens, and continued to be so on later trips to the same region.

Caption: in more recent years, it’s become obvious that Ichthyostega - the classic ‘early tetrapod’ - was not just a formidable, toothy predator, but an unusual, specialised animal with paddle-like hindlimbs, a proportionally short tail and a regionalised vertebral column. Image: Ahlberg et al. (2005).

Was Jarvik right about these animals? Not really, no. The discovery of numerous aquatic specialisations in Ichthyostega and the smaller Acanthostega – published by Mike Coates and Jenny during the early 1990s (Coates & Clack 1990, 1991) – was a huge surprise. Tetrapods didn’t start their history as land-walking animals with pentadactyl hands and feet, as Jarvik (and just about everyone else) had thought, but were aquatic and polydactyl, with 8 fingers and 7 toes (or thereabouts)! Later work showed that Ichthyostega had an unusual ear perhaps specialised for aquatic hearing (Clack et al. 2003) and a remarkable regionalised vertebral column and other peculiarities (Ahlberg et al. 2005) suggesting that, when on land, it might have moved in mudskipper-like fashion (Ahlberg et al. 2005, Pierce et al. 2012, 2013).

Caption: life reconstructions of three early tetrapods worked on by Clack: the big Crassigyrinus, the small, aquatic Acanthostega, and the big (c 1 m long) Ichthyostega. These images are among tens of archaic tetrapods reconstructed for my in-prep textbook. I’ve learnt recently that the Crassigyrinus will soon need revising. Image: Darren Naish.

Jenny’s work wasn’t limited to this fundamental reinterpretation of the earliest tetrapods, but also involved taxa from throughout the Devonian and Carboniferous. She revised knowledge of the remarkable aquatic Carboniferous tetrapod Crassigyrinus (Clack 1998a), described the new species Eucritta melanolimnetes (Clack 1998b), Pederpes finneyae (Clack 2002a, Clack & Finney 2005), Kyrinion martilli (Clack 2003) and others, reported entire new faunal assemblages (Clack et al. 2016) and published key interpretations of such groups as embolomeres (Clack 1987, Clack & Holmes 1988), chroniosuchians (Clack & Klembara 2009, Klembara et al. 2010) and microsaurs (Clack 2011). This extensive, substantial experience made her the right person to publish an authoritative and comprehensive book on Devonian and Carboniferous tetrapods: Gaining Ground: The Origin and Evolution of Tetrapods appeared in 2002 (Clack 2002). It went to second edition in 2012. It’s the best guide to early tetrapod evolution and fills a much-needed gap in the semi-technical literature, and I strongly recommend it to those interested in the subject.

Caption: covers of the first and second editions of Clack’s Gaining Ground. The cover images are, respectively, by Jenny Clack and Julia Molnar. The Clack image shows two individuals engaging in courtship: the green hump at the back left is the body of a second animal.

I never had the privilege of working with Jenny or of accompanying her in the field, but I did have reason to speak to her on several occasions, and to correspond with her. She was friendly and generous with her time and went to trouble to provide me with information and illustrations on an occasion when email wasn’t doing its job. It’s also obvious that she had a sense of humour. Of those new species listed above, Eucritta melanolimnetes translates roughly as ‘creature from the black lagoon’. Some – probably all – of the Ichthyostega and Acanthostega specimens she discovered in Greenland had nicknames: I’m pretty sure I recall seeing that one of them was called Grace Jones. Those who knew her so much better confirm that she was excellent fun, a great leader in the field, and a brilliant mentor.

Caption: several of the archaic Devonian tetrapods study by Clack and her colleagues are excellent, 3D and with very detailed preservation. This image shows an Acanthostega gunnari cast at Musee De L'Histoire Naturelle, Brussels. Image: Ghedoghedo. CC BY-SA 3.0 (original here).

Jenny began her scientific career during the early 1970s and received her PhD at the University of Newcastle Upon Tyne in 1984 under Alec Panchen. Panchen initially offered her a PhD following anatomical discoveries she made while preparing the notoriously difficult holotype specimen of the Carboniferous embolomere Pholiderpeton, initially described by Thomas Huxley in 1869. She joined the University of Cambridge in 1981 and for more than ten years was Assistant Curator at the University’s Museum of Zoology, later being promoted to Senior Assistant Curator and then Curator. She supervised a number of people who went on to become well known in vertebrate palaeontology and evolutionary biology, among them Mike Lee (whose PhD was specifically on pareiasaurs), sauropod expert Paul Upchurch, and actinopterygian fish worker Matt Friedman.

Caption: life-sized models of Ichthyostega and Acanthostega have been made a few times. Here’s the Acanthostega on show at the Sedgwick Museum, Cambridge. I’ve photographed this model several times but none of my photos are that good - I stole this image from Christian Kammerer (source).

Jenny retired in 2015. As you might expect give her scientific achievements, she was highly decorated, holding several palaeontological medals, additional, honorary doctorates and other awards too. Such was mainstream interest in her work and ideas that a 2012 episode of the BBC series Beautiful Minds was devoted to her (it’s viewable here on youtube); among other interesting things, it reveals that she was inspired during her childhood years by newts and freshwater fishes, and that it was learning about Mary Anning which encouraged her to pursue palaeontology. She liked motorbikes and cats, and some of the documentaries that focus on her work show her riding around on her motorbike.

Caption: the Tournaisian rocks of Burnmouth, north of Berwick-upon-Tweed, have, within recent decades, proved an important new locality for tetrapods. As a consequence, Jenny and colleagues set up the TW:eed Project, the acronym standing for Tetrapod World: Early Evolution and Diversity. Jenny (in the green top) stands close to the middle. Image: (c) Rob Clack, used with permission.

My text here only touches on a few aspects of her achievements, life and technical contributions, and I know that much else could be said. I’m very saddened to hear of her passing, very much regarded her as an excellent scientist who published exciting research, and I’ll miss her as a regular attendee of palaeontological meetings. My sincere condolences to her husband Rob, and to those others who knew and loved her.

Some of the text here is adapted from my in-prep giant textbook on the vertebrate fossil record.

• The University of Cambridge website on Professor Clack is here

• An excellent website on Professor Clack’s research and expeditions is here

• The TW:eed Project website is here

Refs - -

Ahlberg, P. E., Clack, J. A. & Blom, H. 2005. The axial skeleton of the Devonian tetrapod Ichthyostega. Nature 437, 137-140.

Clack, J. A. 1987. Two new specimens of Anthracosaurus (Amphibia: Anthracosauria) from the Northumberland coal measure. Palaeontology 30, 15-26.

Clack, J. A. 1988. Pioneers of the land in East Greenland. Geology Today 4 (6), 192-194.

Clack, J. A. 1998a. The Scottish Carboniferous tetrapod Crassigyrinus scoticus (Lydekker) – cranial anatomy and relationships. Transactions of the Royal Society of Edinburgh: Earth Sciences 88, 127-142.

Clack, J. A. 1998b. A new Early Carboniferous tetrapod with a mélange of crown-group characters. Nature 394, 66-69.

Clack, J. A. 2002a. An early tetrapod from 'Romer's Gap'. Nature 418, 72-76.

Clack, J. A. 2002b. Gaining Ground: The Origin and Evolution of Tetrapods. Indiana University Press, Bloomington and Indianapolis.

Clack, J. A. 2003. A new baphetid (stem tetrapod) from the Upper Carbinoferous of Tyne and Wear, U.K., and the evolution of the tetrapod occiput. Canadian Journal of Earth Sciences 40, 483-498.

Clack, J. A. 2011. A new microsaur from the Early Carboniferous (Viséan) of East Kirkton, Scotland, showing soft tissue evidence. Special Papers in Palaeontology 86, 45-55.

Clack, J. A., Ahlberg, P. E., Finney, S. M., Dominguez Alonso, P., Robinson, J. & Ketcham, R. A. 2003. A uniquely specialized ear in a very early tetrapod. Nature 425, 65-69.

Clack, J. A., Bennett, C. E., Carpenter, D. K., Davies, S. J., Fraser, N. C., Kearsey, T. I., Marshall, J. E. A., Millward, D., Otoo, B. K. A., Reeves, E. J., Ross, A. J., Ruta, M., Smithson, K. Z., Smithson, T. R. & Walsh, S. A. 2016. Phylogenetic and environmental context of a Tournaisian tetrapod fauna. Nature Ecology & Evolution 1: 0002.

Clack, J. A. & Finney, S. M. 2005. Pederpes finneyae, an articulated tetrapod from the Tournaisian of western Scotland. Journal of Systematic Palaeontology 2, 311-346.

Clack, J. A. & Holmes, R. 1988. The braincase of the anthracosaur Archeria crassidisca with comments on the interrelationships of primitive tetrapods. Palaeontology 31, 85-107.

Clack, J. A. & Klembara, J. 2009. An articulated specimen of Chroniosaurus dongusensis and the morphology and relationships of the chroniosuchids. Special Papers in Palaeontology 81, 15-42.

Coates, M. I. & Clack, J. A. 1990. Polydactyly in the earliest known tetrapod limbs. Nature 347, 66-69.

Coates, M. I. & Clack, J. A. 1991. Fish-like gills and breathing in the earliest known tetrapod. Nature 352, 234-236.

Klembara, J., Clack, J. A. & Čerňansk‎ý, A. 2010. The anatomy of palate of Chroniosuchus dongusensis (Chroniosuchia, Chroniosuchidae) from the Upper Permian of Russia. Palaeontology 53, 1147-1153.

Norman, D. 1996. [Review of] The Devonian tetrapod Ichthyostega. Palaeontological Newsletter 31, 13-15.

Pierce, S. E., Ahlberg, P. E., Hutchinson, J. R., Molnar, J. L., Sanchez, S., Tafforeau, P. & Clack, J. A. 2013. Vertebral architecture in the earliest stem tetrapods. Nature 494, 226-229.

Pierce, S. E., Clack, J. A. & Hutchinson, J. R. 2012. Three-dimensional limb joint mobility in the early tetrapod Ichthyostega. Nature 486, 523-526.

Ries, C. J. 2007. Inventing ‘the four-legged fish’. The palaeontology, politics and popular interest of the Devonian tetrapod Ichthyostega, 1931-1955. Ideas in History 2, 37-78.

Theropod Dinosaurs of the English Wealden, Some Questions (Part 1)

I have no idea whether I’m known for being a specialist on anything. But of the several zoological subject areas I publish on, among my favourite and most revisited is the dinosaurs of the English Wealden, and in particular the theropods (that is, the predatory dinosaurs) of the English Wealden.

At left: a Wessex Formation scene, depicting Eotyrannus, a compsognathid (at lower right), a pachycephalosaurian Yaverlandia in the middle distance, and the titanosauriform ‘Angloposeidon’. I need to do some new Wealden dinosaur artwork. At right: a younger, slimmer version of this blog’s author, holding the holotype claw of Baryonyx walkeri in 2001 or thereabouts. Images: Darren Naish.

What is the Wealden? It’s a Lower Cretaceous succession – formed of sandstones, siltstones, mudstones, limestones and clays – which was deposited during the Early Cretaceous, its oldest layers being from the Berriasian (and thus about 143 million years old) and its youngest from the early Aptian (and thus about 124 million years old). The sedimentology, subdivisions and terminology of the Wealden are complicated, but all you need to know here is that the whole lot is termed the Wealden Supergroup, that it has an old section called the Hastings Group and a younger section called the Weald Clay Group – both of which crop out on the English mainland – and that there’s also a young section called the Wealden Group that mostly crops out on the Isle of Wight. Finally, you also need to know that the Wealden Group includes the Wessex and Vectis formations. Yikes, even that brief summary was complicated, sorry.

Simplified stratigraphic nomenclature of the Wealden Supergroup. Note that the Hastings Group is much older than the Weald Clay and Wealden groups. The vast majority of Wealden dinosaurs come from the Wessex Formation. Image: Naish (2010).

The really interesting thing about the Wealden is that it’s highly fossiliferous, yielding everything from pollen and diatoms to dinosaurs. Wealden dinosaurs have been hugely important to our evolving understanding of these animals, in part because some of the earliest discoveries – Iguanodon, Hylaeosaurus and Hypsilophodon among them – come from the Wealden succession. Many Wealden dinosaurs have also been famously vexing, in part because they were discovered at a comparatively early stage in our knowledge, in part because their remains have been (and still are) highly incomplete, and in part because their historical taxonomy is a convoluted nightmare. Note also that the circa 20 million year duration of the Wealden means that its dinosaurs were not all contemporaries. Instead, they belonged to a series of distinct faunal assemblages. Within the last few decades, the Wealden has – focusing here on theropods alone – yielded the superstars Baryonyx, Neovenator and Eotyrannus, and its potential to give us really spectacular finds even today is affirmed by additional theropods that are yet to be published.

The Wessex Formation allosauroid Neovenator - here shown with some of its facial bones in partial x-ray - was covered here at TetZoo (ver 3) back in 2017. Our conclusions on the facial anatomy of this dinosaur (Barker et al. 2017) have since been challenged. Image: Darren Naish.

While I could say a whole lot more (I’ve co-authored a whole book on Wealden dinosaurs: Martill & Naish 2001), the point of the article here (and its follow-up, to be published later) is to provide a progress report of sorts on a few contentious or in-prep areas of Wealden theropod research. And I’ll admit right now that the topics I cover here are unashamedly based on my own research interests and projects, sorry. To work…

At left, Martill & Naish (2001) (cover art by Julian Hume). At right, Batten (2011), truly a must-have volume on Wealden palaeontology. Martill & Naish (2001) is now hard to get and only sold at ridiculous prices.

What are you, Yaverlandia? In 1923, Mr F. M.G. Abell discovered the partial skull roof of a fossil reptile at Yaverland on the Isle of Wight. Its thickened bone immediately led Watson (1930) to suggest that it might be from a pachycephalosaur. Fast forward now to the 1970s: Peter Galton – at the time, revising and redescribing all British ornithischians – took this idea and ran with it. He formally named the specimen Yaverlandia bitholus and argued that it was indeed a pachycephalosaur, the most archaic known (Galton 1971). This became the standard take on this dinosaur and the one I supported when writing Dinosaurs of the Isle of Wight in 2001 (Naish & Martill 2001).

At top, the Yaverlandia holotype in (left) ventral and (right) dorsal view. Below, the source of shame. Images: Darren Naish.

During my PhD years I was inspired to think about Yaverlandia anew, mostly because Jim Kirkland and Robert Sullivan (busy at the time with pachycephalosaurs) were pushing the idea that Galton’s identification was very likely wrong. I borrowed the specimen, produced a redescription, had the specimen CT-scanned, and photographed it to death. And I discovered a bunch of new stuff, all of which convinced me that Yaverlandia was not a pachycephalosaur at all, but a theropod. This data formed a chapter of my PhD thesis and brief summaries of my conclusions have been made here and there, including at conferences and in Naish (2011).

Life reconstructions of Yaverlandia are few and far between. This one (seeming to show the animal in a quadrupedal pose: note how the artist has hidden the hand, a classic case of trying to cover up a mistake) is from the fabled Orbis part-work magazine series. I think (but can’t confirm) that the artist was Jim Channel. Image: (c) Orbis.

But the full, detailed explanation of the theropod hypothesis hasn’t yet appeared, though I promise that it will eventually (it’s a work I’m co-authoring with Andrea Cau). As is so often the case with my academic projects, I haven’t been able to make time to finish it (insert reminder about all my academic research being unfunded and done in ‘spare time’: I am not employed in academia). I should also add that a second specimen of Yaverlandia is known and also awaits writing-up. That’s a study I’m doing with Steve Sweetman.

It’s well known in the theropod research community that the full description of this amazing fossil - the holotype of the Spanish ornithomimosaur Pelecanimimus - was done back in the 1990s [UPDATE: nope, 2004], but hasn’t seen print for a bunch of reasons. Consequently, good information on the specimen isn’t (yet) available. Despite that, this photo has been widely shared online. I don’t know who to credit for it.

Are there ostrich dinosaurs in the Wealden or not? Back in the day, I was thrilled by the 1994 description of the remarkable multi-toothed ornithomimosaur Pelecanimimus polyodon from the Barremian Calizas de La Huérguina Formation of Spain. Not just because it’s a neat dinosaur, but because the Calizas de La Huérguina Formation has a lot in common with the Wealden: the two share a list of amphibians, mammals, lizards, crocodyliforms and dinosaurs, this rendering it plausible or likely that Pelecanimimus (or a similar taxon) might await discovery in the Wealden too (Naish et al. 2001, Naish 2002). Predicting the presence of a given group in a given faunal assemblage is a cheap and easy thing to do, and you can award yourself points for prescience and smarts if you’re proved right (even though most people will ignore your prediction), and no-one cares or notices if you never are. So, I’m not looking for a Wealden cookie here. Whatever, “where are the Wealden ornithomimosaurs?” was a question on my mind for a while.

These drawings - produced for Dino Press magazine back in 2002 - look very dated now. They’re supposed to show those smaller theropod groups confirmed for the Wealden (at top) and predicted for the Wealden but still awaiting discovery (at bottom). Images: Darren Naish.

So, I was pretty happy when – in 2014 – Ronan Allain and colleagues announced their discovery of such creatures in the Wealden. They’d discovered a new theropod in the Lower Cretaceous of France (specifically, in the Hauterivian or Barremian of Angeac in Charente, southwestern France) and had used this as a ‘Rosetta Stone’ in the interpretation of other Lower Cretaceous European theropod fossils (Allain et al. 2014). Several Wealden theropods – Valdoraptor, the Calamosaurus tibiae and Thecocoelurus among them – were ornithomimosaurs according to this study (Allain et al. 2014).

In 2014, I superimposed an ornithomimid into the Wessex Formation scene you saw above… this effort was not meant to be entirely serious (and an ornithomimid is the wrong kind of ornithomimosaur anyway). Image: Darren Naish.

I was initially enthusiastic about this proposal and thought that the authors were likely right. But as more and more information has been released on the Angeac theropod, the less like an ornithomimosaur it seems. It looks, instead, like a noasaur. Furthermore, the assorted relevant Wealden remains aren’t as similar to the bones of the Angeac animal as initially argued (Mickey Mortimer pointed this out in an article of 2014). Proper evaluation of what’s going on here will have to wait until a full description of the Angeac theropod appears in print. But if the Angeac theropod is a noasaur, the possibility that it’s close to or congeneric with one or more Wealden theropods remains a likelihood: Thecocoelurus (named for a single cervical vertebra from the Wessex Formation) looks like a noasaur vertebra (Naish 2011)... though that doesn’t necessarily mean that it is (since it also looks like an oviraptorosaur or therizinosaur vertebra in some features).

Mickey produced this image for a 2014 article at The Theropod Database (here).

To bring this round full circle, we might still be missing those predicted Wealden ornithomimosaurs.

Are there other Wealden tyrannosauroids besides Eotyrannus? Loooong-time readers of my stuff – I mean, those who’ve been visiting TetZoo since 2006 – might remember my suggestion from way back that some of the smaller theropod specimens from the Wealden are sufficiently similar to tyrannosauroids from the Lower Cretaceous of China to perhaps be additional small-bodied members of this group. I’m talking about Calamosaurus foxi (named for two cervical vertebrae), Aristosuchus pusillus (named for a partial pelvis and sacrum) and a few additional bits and pieces, including the so-called Calamosaurus tibiae (note the plural there). If these remains do belong to tyrannosauroids, they’re from taxa distinct from Eotyrannus (which everyone agrees is a tyrannosauroid).

The phylogeny I generated for my PhD thesis led me to think that Mirischia might be a tyrannosauroid… in which case Aristosuchus might also be a tyrannosauroid. This isn’t supported, however, in the in-prep Eotyrannus study I’ve co-authored with Andrea Cau. Image: Darren Naish.

I formally suggested a tyrannosauroid identity for Calamosaurus in a 2011 review of Wealden theropods (Naish 2011) but opted to keep Aristosuchus as a compsognathid on account of its similarity with Mirischia from Brazil. However, Mirischia also looks tyrannosauroid-like in some details (it has an anterodorsal concavity on the ilium) and I’ve sometimes wondered if it might also be a member of this clade. Recent results, however, do not support this possibility.

Aristosuchus pusillus is known from a sacrum and its conjoined pubic bones, which possess a notably long, narrow pubic boot (shown in ventral view in the image at bottom right). At left, we see where these bones would fit within the animal (here portrayed as a corpse; the reconstruction is dated and was produced for a conference poster I presented in 1999). Images: Darren Naish, Owen (1876).

So… Calamosaurus, are you a tyrannosauroid or not? When you only have two cervical vertebrae to go on (plus some tibiae that may or may not from the same taxon), it’s about impossible to say, and you can’t resolve things until you have better material. Like, an associated skeleton.

At left, one of the two holotype Calamosaurus foxi vertebrae in multiple views (from Naish et al. 2001). The bone is about 40 mm long in total. At right, a schematic reconstruction showing the two vertebrae in place in the cervical column of a compsognathid- or tyrannosauroid-like coelurosaur (from Naish 2002). Scale bar = 50 mm.

On that note, long-time readers might also recall my mention of a fairly good, associated skeleton of what appears to be a small Wealden tyrannosauroid. But it’s in private hands. I’ve been told by a British palaeontologist that the specimen concerned won’t be available for “this generation” of dinosaur specialists and I should simply forget about it. That’s hard, really hard.

And that’s where we’ll stop now. A second part to this article will be published soon.

Here's your regular reminder that this blog relies on support via patreon, thank you to those providing support already.

For previous TetZoo articles on Wealden theropods and other dinosaurs, see (linking here to wayback machine versions due to destruction or paywalling of everything at versions 2 and 3)…

• Of Becklespinax and Valdoraptor, October 2007

• The world’s most amazing sauropod, November 2007

• Oh no, not another new Wealden theropod!, June 2009

• Concavenator: an incredible allosauroid with a weird sail (or hump)... and proto-feathers?, September 2010

• A truly tiny Cretaceous theropod... from England?, May 2011

• The Wealden Bible: English Wealden Fossils, 2011, November 2011

• Ostrich dinosaurs invade Europe! Or do they?, June 2014 (every archived version of this article lacks the original illustrations, sorry)

• The Sensitive Face of a Big Predatory Dinosaur, June 2017

Refs - -

Allain, R., Vullo, R., Le Loeuff, J. & Tournepiche, J.-F. 2014. European ornithomimosaurs (Dinosauria, Theropoda): an undetected record. Geologica Acta 12, 127-135.

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.

Galton, P. M. 1971. A primitive dome-headed dinosaur (Ornithischia: Pachycephalosauridae) from the Lower Cretaceous of England and the function of the dome of pachycephalosaurids. Journal of Paleontology 45, 40-47.

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.

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

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. 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.

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

Naish, D., Hutt, S. & Martill, D. M. 2001. Saurischian dinosaurs 2: Theropods. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 242-309.

Naish, D. & Martill, D. M. 2001. Boneheads and horned dinosaurs. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 133-146.

Owen, R. 1876. Monograph of the fossil Reptilia of the Wealden and Purbeck Formations. Supplement 7. Crocodilia (Poikilopleuron), Dinosauria (Chondrosteosaurus). Palaeontographical Society Monograph 30, 1-7.

Watson, S. 1930. Cf. Proodon [sic]. Proceedings of the Isle of Wight Natural History and Archaeology Society 1930, 60-61.

Minuscule Hummingbird-Sized Archaic Birds Existed During the Cretaceous

UPDATE (added 13th March 2020): since I published the article below, two relevant matters have come to attention, both of which have implications for the fossil discussed in the article.

Article at left from New Scientist; article at right from New York Times.

The first is that the extraction of amber from the locations concerned is linked with significant humanitarian issues. These make the continued publication and promotion of Burmese amber fossils look unethical; I was only dimly aware of these when writing the article and now regret my (minor) role in the promotion of this discovery (I did plan to delete the article but, on advice, was encouraged to keep it, but add the disclaimer you’re reading now). You can read about the humanitarian issues here, here and here.

Secondly, a number of experts whose opinions I respect have expressed doubts about the claimed theropod status of the fossil discussed below and have argued that it is more likely a non-dinosaurian reptile, perhaps a drepanosaur or lepidosaur (and maybe even a lizard).

A few artists have already produced speculative life reconstructions of Oculudentavis as a lepidosaur or similar reptile. It would have to be a big-brained, shallow-snouted, big-eyed one. Image: (c) Mette Aumala, used with permission.

I did, of course, consider this sort of thing while writing the article but dismissed my doubts because I assumed that - as a Nature paper - the specimen’s identity was thoroughly checked and re-checked by relevant experts before and during the review process, and that any such doubts had been allayed. At the time of writing, this proposed non-dinosaurian status looks likely and a team of Chinese authors, led by Wang Wei, have just released an article arguing for non-dinosaurian status. I don’t know what’s going to happen next, but let’s see. The original, unmodified article follows below the line…

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If you’ve been paying attention to 21st century palaeontological discoveries you’ll know that our understanding of Cretaceous vertebrate diversity has been much enhanced in recent years by the discovery of animals preserved within amber.

A number of really interesting vertebrate fossils in amber have been published in recent years. Among them are the two partial enantiornithine bird wings shown at left (both from Xing et al. 2016a), and the tiny anguimorph lizard Barlochersaurus winhtini (from Daza et al. 2018). Images: Xing et al. (2016), CC BY 4.0, Daza et al. (2018).

These include lizards and snakes (Daza et al. 2016, 2018), a segment of dinosaur tail originally identified as that of a non-bird theropod (Xing et al. 2016b), and various small birds (e.g., Xing et al. 2016a, 2017, 2018, 2019), all of which belong to the archaic, globally distributed group known as the enantiornithines or opposite birds. Today sees the latest of such discoveries, and it’s the most remarkable announced so far. It is – in fact – among the most remarkable of Mesozoic fossils ever announced, and I say this because of the implications it has for our understanding of Mesozoic vertebrate diversity.

Life reconstruction of Oculudentavis khaungraae Xing et al., 2020, depicting it as a tree-dwelling avialan theropod with partly colourful plumage. Image: (c) Gabriel Ugueto, used with permission.

The fossil in question, described in the pages of Nature by Lida Xing, Jingmai O’Connor and colleagues, is the complete, anatomically pristine but minuscule skull of a maniraptoran theropod – specifically, an archaic bird – named Oculudentavis khaungraae (Xing et al. 2020). The skull is preserved in a small amber block (31.5 x 19.5 x 8.5 mm) dating to the Cenomanian age of the Late Cretaceous (making it about 99 million years old). Like virtually all recently described amber vertebrates, it’s from Myanmar (Xing et al. 2020).

One of several images of the tiny Oculudentavis skull provided by Xing et al. (2020), this one (from their Extended Data) showing the specimen in left lateral view. The scale bar is 2 mm. Image: Xing et al. (2020).

When I say that this fossil is ‘minuscule’, I’m not kidding: the entire skull – the whole skull – is 14 mm long (1.4 cm; not a typo)*. This means that – at a very rough guess – the whole animal was around 90 mm (9 cm) long, an estimate I arrived it by producing a very schematic skeleton which equips the animal with a long tail. Xing et al. (2020) very rightly compare Oculudentavis with small hummingbirds: if it had a long bony tail (which it should have, given its inferred phylogenetic position; read on), it would have been longer than the tiny Mellisuga hummingbirds, the total lengths of which are around 50-60 mm, but not by much. It was unbelievably tiny.

* I’m frustrated by the fact that the authors don’t – so far as I can tell – provide the length of the entire skull anywhere in the paper, nor is there a table of measurements or an effort to estimate the animal’s complete size. Which is weird, because surely this is the most interesting thing about it.

A very rough, semi-schematic skeletal reconstruction of Oculudentavis which I produced in order to gain a rough idea of possible size. As you can see, it would have been tiny. The overall form of the skeleton is based on that of jeholornithiform birds; read on. Image: Darren Naish.

The skull of Oculudentavis has a typical ‘birdy’ look. It has a longish, shallow rostrum, large eye sockets, a lot of bone fusion (no, it isn’t a baby) and a rounded cranium where the section posterior to the eyes is short and compact (Xing et al. 2020). The nostrils are retracted, there’s no trace of an antorbital fenestra, the bony bars beneath the eye sockets bow outwards, and there’s a complete bony bar separating each eye socket from the openings at the back of the skull (Xing et al. 2020).

Digital scan of the skull of Oculudentavis in right lateral view (from the Extended Data of Xing et al. 2020). Note the overall toothiness. The dotted lines here show where slices were recorded during the scanning process. Image: Xing et al. (2020).

It’s a toothy little beast, with an atypically high number of conical (or near-conical) teeth lining its jaws all the way back to beneath the eye socket. This is unusual, since the toothrow in toothed birds and bird-like theropods in general normally stops well anterior to the eye. Another unusual feature is that the teeth aren’t located in sockets but are either fused to the jaw bones (the acrodont condition) or located within grooves that extend along the length of the jaws (the pleurodont condition) (Xing et al. 2020). The teeth look prominent, such that it’s hard to understand how they could be sheathed by lip tissue, nor is any such tissue preserved. Remember that beak tissue doesn’t occur in the same part of the jaws as teeth do, so Oculudentavis wouldn’t have had a true horny covering on its jaws. I assume that it had ‘lip’ tissue sheathing its teeth (except perhaps for the tips of the longest ones), as do other terrestrial tetrapods.

Speculative life reconstruction of Oculudentavis, its feathering and other details inspired by Jeholornis and other archaic members of Avialae. I’ve depicted it on the forest floor but am not necessarily saying that this is where it spent all of its time. Image: Darren Naish.

The eyes are directed laterally and the authors note that Oculudentavis likely didn’t have binocular vision (Xing et al. 2020). The sclerotic rings are huge and fill up most of the eye sockets. Xing et al. (2020) use the relative size of the eyes and their sclerotic rings to make inferences about the activity patterns and visual abilities of this animal: they think that Oculudentavis was likely day-active, had relatively small pupils, and perhaps had “unusual visual capabilities”.

The fossil doesn’t just consist of the animal’s bones alone, because synchrotron scanning reveals the presence of a brain (which is about as wide as it is long). Meanwhile, the bony palate preserves traces of its original tissue covering. This is decorated with numerous papillae, the first time such structures have been reported in a fossil theropod (Xing et al. 2020). The authors also refer to a tongue (!!) but it isn’t possible to make this out in the figures they provide, nor do they label it.

Combined, what do these features tell us about the lifestyle and ecology of Oculudentavis? The well-fused skull, prominent teeth and large eyes suggest that this was a predator, presumably of small arthropods. The soft papillae on the palate are of the wrong sort for fish-eating (Xing et al. 2020). Its tiny size and forest habitat imply that it was arboreal or scansorial – as suggested by Gabriel’s artwork above – but the animals that surround it in the cladogram are mostly terrestrial, so the possibility that it foraged in leaf litter or took regular trips to the forest floor are also conceivable, perhaps. Could it have been a predator of worms, molluscs or even tiny vertebrates, like a dinosaurian shrew?

Oculudentavis would have looked noticeably small relative to other Mesozoic birds, though not absurdly so. It’s compared here with Archaeopteryx (at upper left) and an assortment of others, most of which are enantiornithines. These illustrations are for my in-prep giant textbook project. Image: Darren Naish.

What sort of bird is Oculudentavis? For starters, it’s the presence of fused premaxillary and braincase bones, the position and size of the nostril, eye, postorbital region and domed cranium which strongly indicate that Oculudentavis is a member of Avialae, the bird lineage within Maniraptora (though note that the authors prefer the term Aves for said lineage). They included it within a phylogenetic analysis and found it to be one step more crown-ward (meaning, one node on the cladogram closer to living birds) than is Archaeopteryx, which is surprising because it makes Oculudentavis one of the most archaic members of the bird lineage (Xing et al. 2020). This could mean that birds underwent acute miniaturisation almost as soon as they evolved. Several authors – myself and colleagues included (Lee et al. 2016) – have argued beforehand that theropods on the line to birds underwent a gradual and pervasive decrease in size, but we didn’t (and couldn’t) predict that a size decrease of this sort occurred so early in bird history.

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). Image: Davide Bonnadonna.

A World of Tiny Cretaceous Theropods? A key thing here is that we only know about this animal because of its preservation in amber: the rest of the fossil record mostly – the authors suggest – robs us of tiny vertebrates such as this. Could there actually have been many hummingbird-sized miniature theropods of this sort?

Xing et al. (2020) don’t provide a size estimate for Oculudentavis, but they do provide these silhouettes, which show Oculudentavis to scale with a hummingbird and chicken (and part of an ostrich is just visible at far right). Image: Xing et al. (2020).

Here’s where Xing et al.’s (2020) cladogram become especially interesting. The position they propose for Oculudentavis requires that its lineage originated about 150 million years ago, and yet Oculudentavis itself is about 99 million years old. Its lineage, therefore, is at least 50 million years long, in which case there could have been many of these tiny avialan dinosaurs (here, I have to resist the urge to talk about the hypothetical tree-climbing small dinosaurs of Dougal Dixon and George Olshevsky). I emphasise that this speculation assumes that the phylogenetic position Xing et al. (2020) infer is correct; it may not be. Indeed 10% of their trees found Oculudentavis in a different position: within enantiornithines, a possibility which seems ‘more right’ given the identity of other Burmese amber birds. With just a skull to go on, we obviously need more material before we can be especially confident on its phylogenetic position. And on that point, I won’t be surprised if it turns out that Oculudentavis does end up occupying a different position within maniraptoran theropods from the one which Xing et al. (2020) prefer. But none of this affects its minuscule nature, and that’s the real killer point here.

Part of a time-calibrated theropod tree (from Wang & Zhou 2017). According to Xing et al. (2020), Oculudentavis occupies a position more root-ward than Jeholornithiformes, but more crown-ward than Archaeopteryx. If correct, this means that its lineage originated during the latest part of the Jurassic. Image: Wang & Zhou (2017).

What About Other Fossil Vertebrates? If tiny, tiny Cretaceous theropods have remained unknown to us until now, what about other terrestrial vertebrates? I’d always assumed that the truly tiny frogs, lizards and other vertebrates of the modern world – those less than a few centimetres long – were recently evolved novelties of the Cenozoic. But maybe this is completely wrong. Maybe animals of this sort were present in the Mesozoic too, and maybe we’ve missed them due to a size filter which can only be filled by fossils discovered in amber?

The modern world is inhabited by truly tiny lizards and frogs, like this c 3cm SVL Brookesia chameleon and c 1cm Stumpffia frog. Were similarly tiny tetrapods also around in the Cretaceous? Images: (c) Mark D. Scherz, used with permission.

Time will tell. This is really exciting stuff.  

If you enjoyed this article and would like to see me do more, please consider supporting this blog (for as little as $1 per month) at patreon. The more support I receive, the more financially viable this project becomes and the more time and effort I can spend on it. Thank you :)

For previous TetZoo articles relevant to this one, see…

• Bird behaviour, the ‘deep time’ perspective, January 2014

• 50 million years of incredible shrinking theropod dinosaurs, July 2014

• The Romanian Dinosaur Balaur Seems to Be a Flightless Bird, June 2015

• The Most Amazing TetZoo-Themed Discoveries of 2018, December 2018

• A Multi-Species Nesting Assemblage in the Late Cretaceous of Europe, February 2019

Refs - -

Daza, J. D., Bauer, A. M., Stanley, E. L., Bolet, A., Dickson, B. & Losos, J. B. 2018. An enigmatic miniaturized and attenuate whole lizard from the mid-Cretaceous amber of Myanmar. Breviora 563, 1-18.

Daza, J. D., Stanley, E. L., Wagner, P., Bauer, A. M. & Grimaldi, D. A. 2016. Mid-Cretaceous amber fossils illuminate the past diversity of tropical lizards. Science Advances 2 (3), e1501080.

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.

Wang, M. & Zhou, Z. 2017. The evolution of birds with implications from new fossil evidences. In Maina, J. N. (ed) The Biology of the Avian Respiratory System. Springer International Publishing, pp. 1-26.

Xing, L., McKellar, R. C., O’Connor, J. K., Bai, M., Tseng, K. & Chiappe, L. M. 2019. A fully feathered enantiornithine foot and wing fragment preserved in mid-Cretaceous Burmese amber. Scientific Reports 9, 927.

Xing, L., McKellar, R. C., Wang, M., Bai, M., O’Connor, J. K., Benton, M. J., Zhang, J., Wang, Y., Tseng, K., Lockley, M. G., Li, G., Zhang, Z. & Xu, X. 2016a. Mummified precocial bird wings in mid-Cretaceous Burmese amber. Nature Communications 7, 12089.

Xing, L., McKellar, R. C., Xu, X., Li, G., Bai, M., Scott Persons IV, W., Miyashita, T., Benton, M. J., Zhang. J., Wolfe, A. P., Yi, Q., Tseng, K., Ran, H. & Currie, P. J. 2016b. A feathered dinosaur tail with primitive plumage trapped in mid-Cretaceous amber. Current Biology 26, 3352-3360.

Xing, L., O’Connor, J. K., McKellar, R. C., Chiappe, L. M., Bai, M., Tseng, K., Zhang, J., Yang, H., Fang, J. & Li, G. 2018. A flattened enantiornithine in mid-Cretaceous Burmese amber: morphology and preservation. Science Bulletin 63, 235-243.

Xing, L., O’Connor, J. K., McKellar, R. C., Chiappe, L. M., Tseng, K., Li, G. & Bai, M. 2017. A mid-Cretaceous enantiornithine (Aves) hatchling preserved in Burmese amber with unusual plumage. Gondwana Research 49, 264-277.

Xing, L., O’Connor, J. K., Schmitz, L., Chiappe, L. M. McKellar, R. C., Yi, Q. & Li, G. 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature 579, 245-249.

Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays

One of many exciting discoveries made in tetrapod biology in recent decades is that UV-sensitive vision is not just a thing that exists, but a thing that’s widespread.

Caption: 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.

We’ve known since the early 1980s that at least some birds can detect UV wavelengths, and research published more recently has demonstrated its presence in lizards of disparate lineages, in turtles, rodents and, most recently, amphibians. Some of these animals use their UV-sensitive vision to find food (like pollen-rich flowers) and perhaps even to navigate their environments (UV-sensitive vision in certain forest-dwelling birds might enhance their ability to see certain kinds of leaves, for example).

That’s great, but what’s even more surprising – though maybe it shouldn’t be – is that markings and tissue types in some of these animals are visible to other animals with UV-sensitive vision. Furthermore, some tissue types are able to absorb UV and re-emit it within part of the spectrum visible to we humans. It’s this aspect of the UV story – the possibility that UV is absorbed and emitted as visible light (typically blue light) – that we’re talking about hereon, not UV-sensitive vision. Note that the terms used for this phenomenon are slightly contentious among relevant experts. Most agree that the right term is fluorescence whereas others (including my colleague Jamie Dunning) argue that we should use the more specific photoluminescence. I have no proverbial dog in this fight but am going to stick with photoluminescence here seeing as it’s the one we used in the relevant paper.

Caption: in 2018, Jamie Dunning and colleagues reported the discovery of photoluminescence in puffins. Image: (c) Jamie Dunning.

The discovery of photoluminescence in animals is evidently of broad general interest, and I can make this assertion because several recent studies reporting its occurrence have received an unusual amount of public interest. Dunning et al.’s (2018) report on its occurrence in the brightly coloured bill plates of puffins, for example, proved a really popular discovery (Wikinson et al. (2019) followed up with a subsequent study on the keratinous horns of rhinoceros auklets), as did Prötzel et al.’s (2018) discovery of photoluminescent bones in chameleons. Remarkably, Prötzel et al. (2018) were able to show that the ‘glowing’ bones of these lizards are visible through the skin. At the time of writing, a study reporting widespread photoluminescence in living amphibians has just appeared, and it too has received a fair amount of general interest.

Here it’s worth making a critical point on the popularity of these studies in the popular media. There’s no doubt that this stuff is interesting, and certainly of relevance to biologists at large (for one thing, knowing about the distribution of fluorescence/photoluminescence could have all kinds of implications for surveying and collecting). But there’s concern that the studies are being framed in the wrong way, and that more thorough vetting is needed, in places. Also worth noting is that what role photoluminescence actually has to the animals that emit it is controversial, since some workers argue (a) that its visual signalling role hasn’t been sufficiently tested for, and (b) it may simply be too subtle to be of much use to the animals in which it’s present. Keep this in mind when reading the following!

Caption: Prötzel et al.’s (2018) bone-glow research on chameleons shows that the photoluminescing bones of these lizards were actually visible through the skin. Image: David Prötzel.

These caveats notwithstanding, if UV-themed visual displays are widespread in tetrapods, those of us interested in fossil animals are presented with an interesting set of possibilities. We already think that the many extravagant structures of non-bird dinosaurs and pterosaurs – they include cranial horns, crests and casques as well as spikes, spines, sails, bony plates and so on – functioned predominantly in visual display. Could they also have been photoluminescent, and could this have then been used to enhance the display function of the structures in question?

Caption: dinosaurs and pterosaurs are of course notable for their remarkable variety of what I term extravagant structures, a selection of which are depicted here. (a) Parasaurolophus, a hadrosaurid ornithopod. (b) Pachycephalosaurus. (c) Triceratops, a ceratopsid ceratopsian. (d) Dilophosaurus, a theropod. (e) Pteranodon and (f) Tupandactylus the pterodactyloid pterosaurs. (g) Miragaia the stegosaur. (h) Edmontonia the nodosaurid ankylosaur. From Woodruff et al. (2020), images by Darren Naish.

In a brand-new paper published this week in Historical Biology (or on its website, anyway), Cary Woodruff, Jamie Dunning and I set out to consider this very question (Woodruff et al. 2020). At the risk of spoiling the surprise I’ll say that we don’t provide a hard or definitive answer; our aim instead is to bring attention to the possibility that photoluminescence might have been present in some of these animals. We encourage the testing of this possibility and suggest some specific ways in which this testing might be performed. Of incidental interest is that our collaboration evolved from a Twitter discussion (which is currently findable here).

Caption: a palaeontologist ponders new papers on photoluminescence, and then gets talking to one of the relevant researchers. And I chimed in as well, sorry. The rest is history…

I should also add that our idea isn’t especially new. Ever since UV-sensitive vision was first reported in birds back in the 1980s, the idea that extinct dinosaurs might have made use of photoluminescence has been mooted (though, let me make the point again: you don’t need UV-sensitive vision to see photoluminescence). I’ve incorporated photoluminescence into more than one dinosaur-themed media project, most recently Dinosaurs in the Wild.

Caption: the idea that Mesozoic dinosaurs might have been exploiting photoluminescence isn’t altogether new. Here are rough sketches I produced depicting the concept of a photoluminescent Leptoceratops produced for the travelling visitor experience Dinosaurs in the Wild. Image: Darren Naish.

A few specific points are worthy of attention. Above, I mentioned Prötzel et al.’s (2018) chameleon-themed ‘bone glow’ study. Bone-based photoluminescence has also been reported in frogs, specifically in the Brachycephalus pumpkin toadlets (Gouette et al. 2019). Could those dinosaurs superficially similar to chameleons (namely ceratopsians: like some chameleons, they have bony frills and prominent horns) also possess bone-based photoluminescence and, if so, could they exploit it in chameleon-like fashion? Well, probably not, mostly because the much larger size of these dinosaurs means that their skin was too thick for this to work (Woodruff et al. 2020).

Caption: for fun, let’s use toy ceratopsians rather than the real things. Could these dinosaurs have had ‘glowing’ bones as modern chameleons do? No, almost certainly not. Image: Darren Naish.

One of the most unusual things about non-bird dinosaurs possessing extravagant structures is that males and females are extremely similar (albeit not necessarily identical) with respect to the form and proportional size of said structures. As regular TetZoo readers might recall from several articles published here within recent years (see links below), some workers interpret the extravagant structures of Mesozoic dinosaurs as functioning within a model of species recognition. According to this model, the structures function as banners used to signal membership of whatever the respective species is. I don’t think that this is valid for a bunch of reasons and in fact I don’t think that extravagant structures have an important role in species recognition at all (Hone & Naish 2013, Knell et al. 2013). An alternative model posits that extravagant structures mostly have an intraspecific function, work as sociosexual signals of reproductive quality, and evolved within the context of sexual selection. This is the model that I and my colleagues support (Hone et al. 2011, Knell et al. 2012, 2013, Hone & Naish 2013), and a lengthy debate that’s been thrashed out in the literature over the past decade pits species recognition and sexual selection as opposing schools of thought.

Caption: at left, mutual sexual selection at play in the Great crested grebe, as illustrated by Julian Huxley in 1914. At right, cover of the famous issue of TREE which includes Knell et al.’s (2012) seminal review.

But if this is so, why is it that ostensible males and females in the dinosaur species concerned are monomorphic: that is, they have similar extravagant structures? Back in 2011, Dave Hone, Innes Cuthill and I argued that these animals might have evolved their extravagant structures within the context of mutual sexual selection (Hone et al. 2011), this being the strategy where both males and females use their extravagant structures in sociosexual display. But while we know that extant monomorphic animals really are monomorphic, we’re not sure that this is (or was) the case for extinct ones: it could still be that their structures differed in hue, colour or some other visual property. If we’re speculating about the possible presence of photoluminescence in extinct archosaurs, the possibility exists that “monomorphic elaborate structures in pterosaurs and non-bird dinosaurs were not monomorphic in life” but differed in how they photoluminesced (Woodruff et al. 2020, p. 5). We were inspired by the sexually dimorphic photoluminescence of chameleons and Brachycephalus frogs.

Caption: could the in-situ, fully intact armour of ankylosaurs like that of the amazing holotype of Borealopelta, shown here, give insight into the potential of photoluminescence in these animals? Image: CC SA 4.0, original here.

Finally… speculating about the presence of photoluminescence is all very well and good, but can we test for it? In those cases where part of the integument is preserved, we can, by shining blacklights at the respective specimens. The problem, however, is that we might not be seeing the original light-emitting properties of the animal. Seemingly positive results might be a consequence of the fact that various tissues (bone included), minerals and preservatives fluoresce under UV (Woodruff et al. 2020).

As a preliminary test, we looked at the osteoderms of the spectacularly preserved ankylosaurs Borealopelta and Zuul under UV light… we did get results, but it’s difficult to know what, if anything, these results tell us about any condition present in life (Woodruff et al. 2020). I should add that people have been shining blacklights at fossils for a long time and seeing all kinds of interesting results (hat-tip to the pioneering work of Helmut Tischlinger); in no way are we implying that we’re anything like the first to do this.

Caption: people have been examining fossils with UV light for decades. These images show the Jurassic pterosaur Bellubrunnus roethgaengeri, illuminated via the use of UV. Image: Hone et al. 2012 (original here).

And that about wraps things up for now. As will be clear, our paper is not much more than a preliminary set of speculations and suggestions for further work, and isn’t intended to be an in-depth analysis of the proposal. But – as I see it – that’s ok: the scientific literature really shouldn’t be considered focused on results alone, since review, discussion and valid speculation are valuable and worthy too. I hope you agree.

UPDATE (adding 4th March 2020): this article has been somewhat modified relative to its original version, since a misunderstanding on my part meant that I was previously describing photoluminescence as a phenomenon especially relevant to animals with UV-sensitive vision. Substantial thanks to Michael Bok for his interest and assistance and for sending comments which enabled me to modify the article.

This article - and the published technical paper it discusses - happened because of support I receive from a number of excellent and generous patrons. Thank you to that small number of people. Please consider supporting my efforts yourself (for as little as $1 per month), click here.

For previous TetZoo articles on the biology and life appearance of Mesozoic dinosaurs and pterosaurs, see (as usual now, linking to wayback machine versions due to vandalism and paywalling of ver 2 and 3)…

• Zuniceratops and the early acquisition and alleged dimorphism of ceratopsian brow horns, April 2009

• Necks for sex? No thank you, we’re sauropod dinosaurs, May 2011

• Did dinosaurs and pterosaurs practise mutual sexual selection?, January 2012

• Sexual selection in the fossil record, September 2012

• Dinosaurs and their ‘exaggerated structures’: species recognition aids, or sexual display devices?, April 2013

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Dunning, J., Diamond, A. W., Christmas, S. E., Cole, E. L., Holberton, R. L., Jackson, H. J., Kelly, K. G., Brown, D., Rojas Rivera, I. & Hanley, D. 2018. Photoluminescence in the bill of the Atlantic Puffin Fratercula arctica. Bird Study 65 (4), 1-4.

Goutte, S., Mason, M.J., Antoniazzi, M.M., Jared, C., Merle, D., Cazes, L., Toledo, L.F., el-Hafci, H., Pallu, S., Portier, H., Schramm, S., Gueriau, P. & Thoury, M. 2019. Intense bone fluorescence reveals hidden patterns in pumpkin toadlets. Scientific Reports 9, 5388.

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 290, 172-180.

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.

Hone, D. W. E., Tischlinger, H., Frey, E. & Röper, M. 2012. A new non-pterodactyloid pterosaur from the Late Jurassic of Southern Germany. PLoS ONE 7 (7): e39312.

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.

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 28, 250-251.

Prötzel, D., Heß, M., Scherz, M. D., Schwager, M., van’t Padje, A. & Glaw, F. 2018. Widespread bone-based fluorescence in chameleons. Scientific Reports 8, 698.

Wilkinson BP, Johns ME, Warzybok P. 2019. Fluorescent ornamentation in the Rhinoceros auklet Cerorhinca monocerata. Ibis 161, 694-698.

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

Corucia of the Solomon Islands, Most Amazing of Skinks

SKINKS! Again.

A captive Corucia in a commercial collection. Image: Darren Naish.

Skinks are an enormous, globally distributed group of lizards. As of December 2019, there are around 1685 recognised species, accounting for about 25% of living lizard diversity (there are about 6780 lizard species in total), and – perhaps unsurprisingly – I’ve written about them quite a lot at TetZoo… though it’s now hard to appreciate this, since the articles concerned have variously been vandalised, curtailed or paywalled by the hosters of TetZoo ver 2 and ver 3. See links below for the wayback machine versions of these articles.

There’s a lot about skinks in the TetZoo archives, please see the links below. Thank Christ for wayback machine.

Among the most remarkable and striking of skinks is the large Solomon Islands skink or Monkey-tailed skink Corucia zebrata, a prehensile-tailed, mostly green, arboreal skink, and the only member of its genus (though read on). Not only is this amazing lizard green, arboreal and equipped with a powerful prehensile tail, it’s also a giant, especially big specimens reaching 72 cm in total length. This makes it the biggest known skink. It first became known to science in 1856 when indefatigable taxonomist John E. Gray tersely described specimens brought to London by John MacGillivray after his voyage aboard the HMS Herald, the type specimens coming specifically from San Christoval (today termed San Cristobal or Makira) in the Solomon Islands (Gray 1856).

The Solomon Islands. Image by OCHA (original here), CC BY 3.0.

The lizards appear widespread throughout the archipelago (Makira is one of the most southerly islands there) and are variable, differing in eye colour, size, and in the configuration and size of their scales. Some experts think that subspecies should be named to reflect this variation, and the smaller, paler-eyed northern form was named C. z. alfredschmidti in 1997 (Köhler 1997). Maverick Australian bad boy herpetologist Raymond Hoser has claimed the existence of several entirely new species of Corucia, one of which he named for his mother. If you want to know more about Mr Hoser (and why he’s a total joke) see the TetZoo article here.

A captive Corucia in a private collection. Note the dark irides which make this individual look different from some of the other animals shown here. Image: S. Hilgers.

So far, all published work on the phylogeography and variation within Corucia finds it and its divergences to be young; as in, younger than about 4 million years old (Hagen et al. 2012). Yet it must have diverged from its closest living relatives 20 million years ago or more (we can infer this because fossils of other members of the same skink group are this old or older), meaning that the vast bulk of its lineages’ history remains completely unknown, for now.

Gray described Corucia as a new member of the ‘fish-scaled’ skink group. This seems a bit odd today, because we don’t refer to any skink by this moniker (to my knowledge). He evidently regarded it as part of the Australasian skink group that includes Egernia, Tiliqua (the blue-tongues) and kin though. Today we think (on the basis of molecular phylogenetics) that this is correct, and that Corucia is a lygosomine skink (Skinner et al. 2011, Pyron et al. 2013).

Representatives of most (but not all) of the skink lineages currently regarded as ‘families’ by Hedges and colleagues. 1: Mabuya, of Mabuyidae. 2: Acontias, of Acontidae (I think it should really be Acontiidae). 3: Ristella, of Ristellidae. 4: Scincus, of Scincidae. 5: Lygosoma, of Lygosomidae. 6: Egernia, of Egerniidae. 7: Eugongylus, of Eugongylidae. These images are from my in-prep textbook, progress of which can be observed here. Images: Darren Naish.

Traditionally, all skinks are combined in the single family Scincidae. Most herpetologists argue that we should stick with this taxonomic system since there’s no dispute that Scincidae is a clade and thus no real need to shake things up. But some argue that putting all the species of this enormous, complex group into the same single ‘family’ obscures and under-emphasises its diversity and disparity and that it would be more realistic to split it into a whole bunch of families (nine in fact: Acontidae, Atechosauridae, Egerniidae, Eugongylidae, Lygosomidae, Mabuyidae, Ristellidae, Scincidae and Sphenomorphidae) (Hedges & Conn 2012, Hedges 2014). I’ve written about this situation before: see the articles below for more. If we follow this revised family-level classification, Corucia is part of Egerniidae.

Substantially simplified cladogram depicting lygosomine skink phylogeny, mostly based on Pyron et al. (2013). Images (top to bottom): Wolfgang Wuster, H. Zell, $Mathe94$, Benjamint444 (all CC BY-SA 3.0), Mark Stevens (CC BY 2.0), W. A. Djatmiko, S. Caut et al. (both CC BY-SA 3.0).

The name Corucia is derived from ‘coruscus’ (meaning shimmering, and referring to the shiny scales), while zebrata is a reference to the stripes present in the specimens Gray was familiar with. Given that Solomon Islanders know this lizard and eat it, there was and is surely indigenous knowledge of the species and probably lore about it, though I haven’t encountered such so far. It’s generic name shouldn’t be confused with that of the Cretaceous fossil lizard Carusia, a possible relative of the living xenosaurids.

Here in the UK, it’s currently not difficult to encounter Corucia in captivity. I should add that it does well if conditions are right: as a canopy-dwelling lizard it needs tall branches with suitable retreats, and some collections (most notably the Philadelphia Zoo) have been breeding Corucia for over 40 years now. They’re not especially active during peak visitor time at zoos, mostly because they’re crepuscular. They’re also exclusively herbivorous and are in fact the only skinks said to be committed to a plant diet. Leaves, shoots, flowers and fruit are all consumed, including those of toxic species. Their dung has a distinctive aroma and it’s apparently possible to locate trees inhabited by this species by smell alone: Harmon (2002) used this technique, making his study “the first documented use of olfactory cues to locate skinks in the wild” (p. 177).

Fine side-eye from this captive Corucia at Bristol Zoo, UK. Image: Darren Naish.

Corucia is viviparous with a 6 to 8 month gestation, but the big deal about its viviparous strategy is that its babies are proportionally enormous, being about half the size of the mother. They can be over 30 cm long and weigh 175 g. Unsurprisingly, only a single baby is normally produced, though rare cases of twins and triplets are on record.

Corucia is also a social skink. In this, it’s far from unique, since egerniids of more than 20 species live together in family groups and even exhibit monogamy, kin recognition, colonial living and co-operation. Juvenile Corucia sometimes stay with their parental group for an extended period and mothers are reported to be highly protective of newborn juveniles (Wright 2007), which is what theory predicts given the substantial material investment involved in growing such a large baby. Also interesting is that not all the adults which form social groups in this species appear related (Wright 1996), and that Corucia groups are even known to allow orphaned juveniles to join their groups (read on…). Some juveniles do apparently leave their parental group to join others (Wright 2007).

Dark-eyed captive Corucia, and here’s proof that this arboreal lizard will - in captivity - drink from standing water (at least some arboreal lizards don’t do this, they rely only on water droplets on leaves). Image: S. Hilger.

Studies of wild-living Corucia on Ugi Island in the Solomon archipelago showed that individuals living less than 150 m apart were likely to be related, but also that individuals wandered for several kilometres (Hagen et al. 2013). Telemetry results obtained in an earlier study (Hagen 2011) indicate that this sort of dispersal is unusual, however, given that Corucia is mostly sedentary with home ranges being equivalent to the canopy of a single tree. Maybe this explains why groups are apparently happy to ‘adopt’ lone youngsters – they may well be related to the members of the group already. After all, we know that kin selection is at play elsewhere in social egerniids.

One of the latest papers discussing social behaviour in these skinks is also among the most shocking, since it reports the occurrence of a Corucia group living together in a deep tree hole, and one that was flooded at its bottom. Remarkably, some of the Corucia in the hole were fully submerged and located beneath the water surface at the time of discovery. To my supreme frustration, I can’t locate this publication right now, even though I recall downloading it (it was a short note in, perhaps, Salamandra or Journal of Herpetology). Let me know if you know the paper concerned. It was such a bizarre report that more information is needed. And I guarantee that it’s legit and that I didn’t dream it.

A captive Corucia at Bristol Zoo. Note the sharply curved claws and interesting nose in these lizards. Image: Darren Naish.

Finally, what does the future hold for this amazing lizard? Unsustainable destruction of forests on the Solomon Islands poses a problem, as does local hunting for the pot and collecting for the pet trade: between 1992 and 1995, 12000 animals were exported for this reason, mostly to the USA (Mann & Meek 2004). Consequently, Corucia is now being considered for inclusion on Appendix I of CITES, with captive breeding likely being crucial to its persistence.

Another captive Corucia. This image is useful and interesting because it shows the cross-sectional shape of the body: note that the side of the body is flat and that there’s an obvious change in angle between the side and dorsal surface. Image: TimVickers (original here), public domain.

A giant, fully herbivorous, slow-breeding, social skink is such a special animal that we must make effort to ensure its survival into the future. And that’s where we must end.

If you enjoyed this article and would like to see me do more, please consider supporting this blog (for as little as $1 per month) at patreon. The more support I receive, the more financially viable this project becomes and the more time and effort I can spend on it. Thank you :)

For previous TetZoo articles on skinks, see…

• Mystery emo skinks of Tonga!, October 2010

• Isopachys: worm-like skinks from Thailand and Myanmar, November 2010

• Hammer-toothed skink SMASH!, November 2012

• Skinks, Skinks, Skinks!, October 2014

• Terror skinks, social skinks, crocodile skinks, monkey-tailed skinks… it's about skinks (skinks part II), October 2014

• Sandfishes and kin: of sand-swimming, placentation, and limb and digit reduction (skinks part III), November 2014

• The Madagascan Skink Amphiglossus Eats Crabs, February 2016

Refs - -

Gray, J. E. 1856. New genus of fish-scaled lizards (Scissosarae) from New Guinea. Annals and Magazine of Natural History (2) 18: 345-346.

Hagen, I. J. 2011. Home ranges in the trees: radiotelemetry of the Prehensile tailed skink, Corucia zebrata. Journal of Herpetology 45, 36-39.

Hagen, I. J., Donnellan, S. C. & Bull, C. M. 2012. Phylogeography of the prehensile-tailed skink Corucia zebrata on the Solomon Archipelago. Ecology and Evolution 2, 1220-1234.

Hagen, I. J., Herfindal, I., Donnellan, S. C. & Bull, C. M. 2013. Fine scale genetic structure in a population of the prehensile tailed skink, Corucia zebrata. Journal of Herpetology 47, 308-313.

Harmon, L. J. 2002. Some observations of the natural history of the Prehensile-tailed skink, Corucia zebrata, in the Solomon Islands. Herpetological Review 33, 177-179.

Hedges, S. B. 2014. The high-level classification of skinks (Reptilia, Squamata, Scincomorpha). Zootaxa 3765, 317-338.

Hedges, S. B . & Conn, C. E. 2012. A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae). Zootaxa 3288, 1-244.

Köhler, G. 1997. Eine neue Unterart des Wickelschwanzskinkes Corucia zebrata von Bougainvillle, Papua-Neuguinea. Salamandra 33 (1), 61-68.

Mann, S. L. & Meek, R. 2004. Understanding the relationship between body temperatureand activity patterns in the giant Solomon Island skink, Corucia zebrata, as a contribution to the effectiveness of captive breeding programmes. Applied Herpetology 1, 287-298.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 2013, 13:93.

Skinner, A., Hugall, A. F. & Hutchinson, M. N. 2011. Lygosomine phylogeny and the origins of Australian scincid lizards. Journal of Biogeography 38, 1044-1058.

Wright, K. 1996. The Solomon Islands skink. Reptile & Amphibian Magazine 3 (2), 10-19.

Wright, K. M. 2007. Captivating giants. Reptiles Magazine 15 (12), 54-68.

Beautiful, Big, Bold Dinosaur Books: of Molina-Pérez and Larramendi’s Theropods, Rey’s Extreme Dinosaurs 2, and Parker et al.’s Saurian

One of the reasons you read TetZoo is because of the dinosaurs, and among the dinosaur-themed things I write about on fairly regular basis are new(ish) dinosaur-themed books.

Partly because I’m way overdue on the book reviews I planned to write during 2019, I’m here going to talk about some recently-ish published dinosaur-themed books that you’d do well to buy and read, if you wish, or can. I’ve written about recently-ish published dinosaur-themed books on quite a few recent occasions; see the links below for more. Let’s get to it.

Encyclopedia of Dinosaurs: the Theropods, by Rubén Molina-Pérez and Asier Larramendi

I can say right out of the gate that this 2019 work is one of the most spectacular dinosaur-themed works that has ever seen print. Think about that for a minute, since it’s a pretty grand claim. Yes. This book is spectacular: big (288 pages, and 24.5 cm x 30 cm), of extremely high standard, packed with information, and containing a vast number of excellent and highly accurate colour life reconstructions. Originally published in Spanish, it has now been translated (by David Connolly and Gonzalo Ángel Ramírez Cruz) and published in English by London’s Natural History Museum. The book consists of eight sections, which variously go through the theropod cladogram, discuss geographical regions and the theropods associated with them, and review theropod anatomy, eggs, footprints and so on. And it’s packed with excellent illustrations… hundreds of them.

A selection of pages from Molina-Pérez & Larramendi (2019). At left, eggs depicted to scale (with a basketball). At right, just two of the many pages that feature theropod skeletal elements. Images: Molina-Pérez & Larramendi (2019).

The art is great – the majority of colour images being by the phenomenally good and reliable Andrey Atuchin – and I’d recommend that anyone interested in the life appearance of dinosaurs obtain the book for its art alone. I have one criticism of the art though, which is that the colour schemes and patterns used for some of the animals are occasionally based on those of living animals (most typically birds).

Just two of the many UNNAMED theropod species reconstructed in the book. Exciting stuff! The humans that feature in the book are an interesting lot. Images: Molina-Pérez & Larramendi (2019).

Familiar theropods of many sorts are illustrated, but a major plus point is that many of the animals depicted are either obscure and mostly new to the world of palaeoartistic depictions (examples: Spinostropheus, Dryptosauroides, Bonapartenykus, Nanantius, Gargantuavis) or are as-yet-unnamed species: animals which clearly represent something new (since they’re the only member of their group known from the relevant geographical region and segment of geological time) but have only been referred to by their specimen numbers or by a ‘cf’ attribution (a theropod called, for example ‘cf Velociraptor mongoliensis’ is being compared by its describers to V. mongoliensis and is clearly very much like V. mongoliensis, but quite possibly not part of the species and perhaps something new. The ‘cf’ is short for ‘confere’, as in: compare with).

If the reconstructions in the book are anything to go by, Spinostropheus - according to one specimen (an ulna) it could reach huge sizes - was among the most remarkable of theropods. Just look at it. Image: Molina-Pérez & Larramendi (2019).

Two issues strike me as problematic though. One is that the arrangement is really difficult to get to grips with, and it’s taken me numerous attempts to understand and appreciate why the book is arranged the way it is. The second issue concerns what appears to be a suspiciously high degree of taxonomic precision for footprints. The authors depict footprints, said to be representative of the different theropod groups covered in the taxonomy section at the start of the book, and seem confident that the tracks concerned (which are often fairly nondescript) were made by species belonging to the relevant group. It’s hard to be convinced that this is reliable, except in a very few cases: I’m happy to agree that the giant Tyrannosauripus pillmorei track, for example, really was made by a member of Tyrannosauridae. Then again, maybe the authors have devised a new track identification method that isn’t yet known to the rest of us.

The several montages in the book are truly things of beauty. Image: Molina-Pérez & Larramendi (2019).

And on that note, it’s obvious that a vast quantity of novel science was performed as part of the background research for this book. The methods and data used by the authors are explained up front. Impressive stuff, and stuff which should be published in the technical literature at some point. Are their results always ‘good’? Well, I have my misgivings about the idea that Dinornis the moa was the fastest (non-flying) theropod ever and capable of sprinting at a phenomenal 81 km/h…

So, so many diagrams of tracks and trackways. Image: Molina-Pérez & Larramendi (2019).

Despite my minor misgivings, this book is attractive enough and interesting enough that it’s a must-have for those seriously interested in dinosaurs and in artistic depictions of them. Buy it if you can. This volume promises to be the first in a series. At the time of writing, the second of these – devoted to sauropods – is being advertised and is due to appear soon. Given that there’s every reason to assume that its artwork and overall quality will be similar to that of the theropod volume reviewed here, I have very high hopes and look forward to seeing it.

Molina-Pérez, R. & Larramendi, A. 2019. Encyclopedia of Dinosaurs: the Theropods. The Natural History Museum, London. pp. 288. ISBN 9780565094973. Hardback. Here at publishers. Here at amazon. Here at amazon.co.uk.

Luis Rey’s Extreme Dinosaurs 2: the Projects

Luis Rey (who blogs here) has been active in the palaeoart world for a few decades now, and most people interested in the portrayal of dinosaurs in art will be familiar with his vibrant, bold and dynamic style. Luis’s work has appeared in museum installations, exhibitions, and in numerous publications, including books. Most notable among these are his own Extreme Dinosaurs (Rey 2001) and The Big Golden Book of Dinosaurs, by Robert Bakker (Bakker 2013). I feel it would be wrong at this point to avoid mentioning the fact that Luis and I were regular correspondents back when the first of those books appeared, but that my dislike of the 2013 book – which I made clear in a TetZoo review – coincided with a cessation in any contact we used to have. But things have moved on; Luis’s art has continued to evolve and let’s put all of that behind us.

A collection of Rey works at TetZoo Towers. Image: Darren Naish.

Extreme Dinosaurs 2: the Projects discusses the intellectual and artistic background to several dinosaur-themed museum installations which Luis has created, but does so in an evolutionary fashion such that they’re used to describe our improving knowledge of the Mesozoic world. Luis was illustrating colourful, fully feathered dromaeosaurs, oviraptorosaurs and so on at a time when the majority of relevant academics were dead against this, so it would be fair to see him as one of several artists who were predicting things that would prove correct in the end. I was on side too, and consequently was a vociferous Rey advocate in the early part of my academic career, deliberately using his reconstructions of dromaeosaurs – even the turkey-wattled, shaggily feathered ones – in conference presentations and publications. And for all the success of All Yesterdays and its associated movement, we have never forgotten that Luis was saying many of the same things already.

Some representative pages from Extreme Dinosaurs 2. At right, note the person wearing an oviraptorosaur costume while sat in a nest.

Maniraptorans, thyreophorans, ceratopsians, dinosaur eggs and nesting and the dinosaurs (and other Mesozoic reptiles) of Mexico all get coverage here as Luis explains the thinking behind new pieces of art and also how and why he’s modified older ones. Our understanding of feather arrangements in non-bird maniraptorans have improved a lot in recent years, plus we have so much new data on ankylosaur armour, ceratopsian skin and so on. If you haven’t been keeping up, this book would be a good primer. The text is concise and written in a friendly, informal style.

More representative pages, this time depicting therizinosaurs. Brightly coloured faces, bold patterns on the feathering, inflatable throat structures… what’s not to love? Image: Rey (2019).

I’m still not keen on the photo-bashing that’s now integral to the Rey style and don’t find it effective or successful. A few pieces included in the book don’t, therefore, work for me (examples: Santonian hadrosaur hassled by theropods on pp. 104-5, the Labocania scene on pp. 106-7). But can I please emphasise that I still find this a valuable book, and I very much recommend it as an interesting addition to the palaeoart stable, the autobiographical angle in particular being useful. My copy of Extreme Dinosaurs 2: the Projects is an advance softback version but I understand that a hardback is available too.

Rey, L. 2019. Extreme Dinosaurs 2: the Projects. Imagine Publishing, London/Metepec. pp. 139. ISBN 978-0-9933866-2-6. Softback/Hardback. Here at amazon.co.uk.

Parker et al.’s Saurian: A Field Guide to Hell Creek

Even if – like me – you’re not a video game buff and have little interest in video games or the playing of them, chances are high that you’ve heard about Saurian, a role-playing, survival simulation experience in which you play the role of a dinosaur negotiating a Maastrichtian environment modelled on that of the Hell Creek Formation. You have to avoid predators, find and procure food, raise babies, and live as long as possible. I’ve played The Simms, back in the day, and recall the urge to sink hours of time into living a virtual tiny life, so I can understand the appeal.

Representative pages, here showing the raptor prey restraint model in action. Poor pachycephalosaur. Image: Parker et al. (2019).

The relevance of Saurian to the TetZooniverse is that the game has been designed and built according to an incredibly high scientific standard, the team behind it having done a vast quantity of research on everything relevant to the Hell Creek world. And the good news is that this work hasn’t been wasted. This book – written by Tom Parker, featuring the artwork of Chris Masna and RJ Palmer (of Detective Pikachu fame and much else), and produced following consultation with a long list of relevant palaeontologists – is the result: it includes a vast quantity of amazing concept art, detailed vignettes and scaled artwork of the organisms and environments that feature in the game, and is simply a joy to look at. The text is brief but functions well. The book is a sturdy softback (20 x 24 cm) of 182 pages, printed to excellent, glossy standard.

RJ Palmer’s T. rex is one of the stars of the show. Some of you will know that the Saurian team abandoned an earlier, more feathery version. Assorted humans function as scale bars; you can likely guess who this one is based on. Image: Parker et al. (2019).

As should be clear by now, the thoroughness of the project’s world-building means that there’s more information here on Maastrichtian North American trees, rainfall patterns, swamps, beaches, amphibians, lizards, crocodyliforms, fishes and so on than you’ve ever seen before. The result is one of the most interesting, detailed and attractive volumes dedicated to Late Cretaceous life. This book is a must-have for those seriously interested in palaeoart and in seeing prehistoric animals and environments depicted in detail, but it’s also good enough that those with a scientific or technical interest in Late Cretaceous life should obtain it too.

Oh wow, so much of the art in this book is just phenomenal. This scene depicts competition among scavengers at a carcass. You might just be able to see the anguimorph lizard inside the body cavity. Image: Parker et al. (2019).

Parker, T., Masna, C. & Palmer, R. J. 2019. Saurian: A Field Guide to Hell Creek. Urvogel Games. pp. 182. Softback. Here at publishers [BUT CURRENTLY OUT OF STOCK].

That’s where we’ll end for now. A few more dinosaur-themed book review are due to appear here soon, including of Donald Prothero’s The History of Dinosaurs in 25 Fossils and Michael Benton’s The Dinosaurs Discovered: How a Scientific Revolution is Rewriting Their Story.

If you enjoyed this article and would like to see me do more, please consider supporting this blog (for as little as $1 per month) at patreon. The more support I receive, the more financially viable this project becomes and the more time and effort I can spend on it. Thank you :)

For previous TetZoo dinosaur-themed book reviews, see… (linking here to wayback machine versions due to paywalling and vandalism of TetZoo ver 2 and 3 articles)…

• Long and Schouten's Feathered Dinosaurs, a review, January 2008

• Luis Chiappe's Glorified Dinosaurs: The Origin and Early Evolution of Birds, January 2011

• What they’re saying about The Great Dinosaur Discoveries, November 2011

• Greg Paul’s Dinosaurs: A Field Guide, August 2012

• New Dinosaur Books, Part 1: Barrett on Stegosaurs, August 2018

• New Dinosaur Books, Part 2: Ben Garrod’s ‘So You Think You Know About… Dinosaurs’ Series, August 2018

• The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved, November 2011

Refs - -

Bakker, R. T. 2013. The Big Golden Book of Dinosaurs. Random House, New York.

Rey, L. 2001. Extreme Dinosaurs. Chronicle Books, San Francisco.

The Mostly Arboreal African Gastropholis Lizards

TetZoo hasn’t had nearly enough lizards lately. If you’re a regular reader you might recall the slow-burn series on LACERTIDS, the Old World group that are the ‘typical’ lizards of western and central Europe and hence the first group to become known to European scientists.

Caption: if you’re European, lacertids are the lizards you know best. In the UK, we only have two natives, one of which is Zootoca vivipara, the Viviparous lizard. Here’s a wild individual. Image: Darren Naish.

European lacertids – thinking here of Zootoca (viviparous lizards), Lacerta (sand lizards, green lizards and kin) and Podarcis (wall lizards) in particular – are quite well known and it’s easy to find information on them; if, that is, you’ve combed through books on lizards in quest of hot lacertid info. But there are a whole bunch of African taxa within Lacertidae: what’s the deal with them, and why don’t we hear more about them? Well, comparatively little is known about them, good photos of them aren’t that abundant, and they’re rare in museum collections (Arnold 2004).

Caption: a captive Green keel-bellied lizard Gastropholis prasina, in repose. Check out that tail! Images: Darren Naish.

I recently got to see a live lacertid of a type I’d never seen before: a Green keel-bellied lizard Gastropholis prasina, a shockingly long-tailed, green, east African lacertid endemic to the coastal forests of Mozambique, Tanzania and Kenya. It reaches 40 cm in total (which seems pretty huge for a lacertid), but is more usually 25-35 cm. Most of this is formed by the slender, prehensile tail. These are diurnal lizards, and one of several things that makes them unusual among lacertids is that they’re arboreal, and do just about everything up in the trees. They sleep there, fight there, mate there and lay their eggs there (in damp hollows). Individuals of G. prasina have been found 12 metres up in trees (Spawls et al. 2018).

Caption: an 1886 illustration of the Striped keel-bellied lizard G. vittata, from J. G. Fischer’s publication. Image: J. G. Fischer, public domain.

Well…. the Green keel-bellied lizard lives this way, anyway. Animals refuse to be constrained by anatomy and not all members of this genus are arboreal. The Striped keel-bellied lizard G. vittata is apparently terrestrial despite being proportionally pretty much exactly like G. prasina and being equipped with a really long, prehensile tail (Spawls et al. 2018). What this most likely shows is that G. vittata has only recently transitioned to terrestrial life, and that its anatomy still harks back to an arboreal ancestry.

Caption: you can see why this species is called Gastropholis echinata (the species name meaning ‘spiny’). It’s from a 1919 description of the species by K. P. Schmidt. Image: K. P. Schmidt/AMNH, public domain.

Two other species are presently included in Gastropholis. G. echinata occurs throughout those countries bordering the Gulf of Guinea while G. tropidopholis is endemic to the Democratic Republic of the Congo.

Caption: another Gastropholis species: the Congolese G. tropidopholis. Again this image is from a 1919 paper by K. P. Schmidt. Image: K. P. Schmidt/AMNH, public domain.

Convergence with grass lizards…. and monitors? As implied by the name, one of the unusual things about these lizards is that their ventral scales are keeled. That’s pretty odd, since lizards (and other reptiles) ‘ordinarily’ have smooth ventral scales. Keeled ventral scales are also present in at least some species of Adolfus – another climbing African lacertid – and also in Takydromus, the east Asian grass lizards. Like Gastropholis, Takydromus has a strikingly long tail which it uses in climbing (albeit in grasses rather than high in trees).

These lizards also (together with Philochortus, another poorly known African lacertid) share especially tall, blade-like neural spines on some of their caudal vertebrae (Arnold 1997, 2004). Neither Takydromus nor Philochortus are at all close to Gastropholis in lacertid phylogeny, so any similarities have to be a product of convergence (Arnold 2004). If you want to learn more about Takydromus, I wrote about it here at ver 2, back in 2013.

Caption: a Takydromus - an Asian grass lizard - in captivity. These lacertids are similar to the African Gastropholis species in several respects, but are not that closely related to them. Image: Darren Naish.

On the subject of convergence, I can’t help but also regard these mostly arboreal, often green lacertids as convergent with the also long-tailed, also green prasinoid tree monitors of northern Australasia. Prasinoids – the Green tree monitor Varanus prasinus and its several close relatives – are larger than Gastropholis species (40 cm vs 60-110 cm, total length) and have several varanid-flavoured features lacking in lacertids (a long forked tongue, blade-like teeth, longer and more dextrous digits and – probably – enhanced cognition). Prasinoids seem more formidable as predators than is Gastropholis, since they predate on small mammals, big spiny insects and so on. This could just be a consequence of size though, and Gastropholis is reported to predate on smaller lizards so might well be quite formidable too (Spawls et al. 2018). If you’re wondering why there are both Varanus and Gastropholis taxa within the species name prasinus (or prasina), it’s because this means ‘light green’.

Caption: I’ve had reason to draw prasinoids - green tree monitors - on several occasions, and here’s a montage depicting several species. Can these varanids be considered convergent with the long-tailed, arboreal lacertid Gastropholis? Hmm, maybe. Images: Darren Naish.

I’ve written about prasinoids at TetZoo: please go here (reminder that I now link to wayback machine versions of my ver 3 articles, since SciAm has either mutilated my articles or paywalled them).

Where in the family tree? Where do the Gastropholis species fit within the lacertid radiation? None of these lizards are well known and they tend to be missed entirely from popular works that mention or discuss lacertids, this creating the impression that Lacertidae doesn’t really have a presence in the African tropics, which it totally does. Within recent decades it’s been widely agreed that these lizards are part of the equatorial lacertid clade that includes Adolfus and kin. This whole group is probably part of Eremiadini, the lacertid clade that also includes the fringe-toed lizards (Acanthodactylus) and racerunners (Eremias) (e.g., Peréz I de Lanuza & Font 2014)… though be sure to see the TetZoo article on racerunners and kin to get some perspective on the different phylogenetic proposals that have been put forwards for these animals.

Caption: a substantially simplified lacertid phylogeny, showing the approximate structure pieced together in assorted studies. Gastropholis by Darren Naish, Acanthodactylus by Richard Hing, Eremias by Yuriy75 (CC BY-SA 3.0; original here), Takydromus by Acapella (CC BY-SA 3.0; original here), Lacerta by Darren Naish, Gallotia by Petermann (CC BY-SA 3.0; original here), Psammodromus by Wolfgang Wüster. Image CC BY-SA.

Arnold (1989) examined anatomical features and found Gastropholis to form a clade with Bedriagaia, Holaspis and Adolfus, the four forming what he termed ‘the Equatorial African clade’. I have to mention in passing that Holaspis is especially interesting because it’s able to glide and has seemingly evolved this ability as an exaptation (Arnold 2002). That’s a story for another time though. Greenbaum et al. (2011), using data from several genes, found Gastropholis to be the sister-group to Adolfus sensu stricto (Adolfus of tradition is paraphyletic with respect to Holaspis, so some of its populations are now separated and in the 2011 genus Congolacerta).

A note on the future. What else do we know about these lizards? They’re suspected to be in decline, given that the forests they inhabit are being destroyed, but hints that they might be adaptable enough to persist come from the fact that they’ve moved into cash crop plantations in some places (Spawls et al. 2018). The individual shown in my photos above and below is a captive and part of the pet trade. The trade in exotic reptiles (and amphibians) is mostly evil, exploitative and the opposite of anything that can be considered sustainable and beneficial. Having said that, the fact that populations of animals endangered in the wild are now being bred in captivity might provide some kind of safeguard for the future. That’s where we’ll end for now, but we’ll return to lacertids - and other lizards - sometime in the near future.

Caption: Green keel-bellied lizard in profile. Image: Darren Naish.

For previous TetZoo articles on lacertids (linking here to wayback machine versions to avoid issues with the hosting sites, SciAm in particular), see…

• The New Forest Reptile Centre, August 2012

• It’s high time you were told about Psammodromus, July 2013

• Tale of the Takydromus, September 2013

• Racerunner Lizards of the World Unite, October 2014

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Arnold, E. N. 1997. Interrelationships and evolution of the east Asian grass lizards, Takydromus (Squamata: Lacertidae). Zoological Journal of the Linnean Society 119, 267-296.

Arnold, E. N. 1989. Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. Bulletin of British Museum of Natural History (Zoology) 55, 209-257.

Arnold, E. N. 2002. Holaspis, a lizard that glided by accident: mosaics of cooption and adaptation in a tropical forest lacertid (Reptilia, Lacertidae). Bulletin of British Museum of Natural History (Zoology) 68, 155-163.

Arnold, E. N. 2004. Overview of morphological evolution and radiation in the Lacertidae. In Pérez-Mellado, V., Riera, N. & Perera, A. (eds) The Biology of Lacertid Lizards. Evolutionary and Ecological Perspectives. Institut Menorquí d’Estudis. Recerca 8, 11-36.

Greenbaum, E., Villanueva, C. O., Kusamba, C., Aristote, M. M. & Branch, W. R. 2011. A molecular phylogeny of equatorial African Lacertidae, with the description of a new genus and species from the Democratic Republic of the Congo. Zoological Journal of the Linnean Society 163, 913-942.

Peréz I de Lanuza, G. & Font, E. 2014. Ultraviolet vision in lacertid lizards: evidence from retinal structure, eye transmittance, SWS1 visual pigment genes and behaviour. The Journal of Experimental Biology 217, 2899-2909.

Spawls, S., Howell, K., Hinkel, H. & Menegon, M. 2018. Field Guide to East African Reptiles, 2nd Edition. Bloomsbury, London.

TetZoo Reviews Zoos: the Isle of Wight Zoo

Regular readers might be aware of my ‘TetZoo Reviews Zoos’ series which hasn’t – so far – been all that active or successful when it comes to completeness of coverage or regularity of appearance. I aim to rectify that through the course of this year.

Caption: there are a few articles in the TetZoo archives on zoos, but not nearly as many as I’d hoped there’d be by now.

Today, I want to talk about a small zoo that’s close to me here on the southern coast of the UK: namely, the Isle of Wight Zoo at Sandown, Isle of Wight. The zoo has a very good website, and much of the information relayed here is derived from it.

Caption: the zoo’s website is here. It has a facebook page here.

Sandown is on the island’s south-west coast; the zoo itself is actually adjacent to the seafront and award-winning beach there and is just a few minutes down the road from Dinosaur Isle Museum. Indeed, it’s close enough to the sea for it to be in danger given the sea level rise that’s due to happen over the next few decades. Like so many zoos, the Isle of Wight Zoo started out as a small, family-run facility essentially created for profit and entertainment. It has more recently (2017) made the transition to being a conservation-oriented charity – termed The Wildheart Trust – devoted to education and animal rescue, and with ties to conservation projects in India, Madagascar and the UK.

Caption: it was wet, cold and misty when Will and I visited. Image: Darren Naish.

The zoo specialises in particular as a refuge for big cats – mostly tigers – who’ve been rescued from sad treatment and neglect at circuses, indeed I think it’s the UK’s largest collection of tigers. Most of the zoo’s tigers suffer from some degree of lameness or injury due to their life in inappropriate conditions and the zoo and its vets have done much to improve their quality of life. The bloodlines of the zoo’s tigers are unknown and there have never been any plans to breed them.

Caption: one of the zoo’s several tigers. Some of the tigers here have lived into their 20s, which makes them among the oldest tigers in the world. The world’s oldest tigers have variously been 22, 25 or similar. Image: Darren Naish.

The zoo’s surrounds are interesting in that it’s constructed within the ruins of a Victorian fort, the big concrete blocks that form the upper parts of its outer walls being distinctive parts of its architecture. A tiger statue, perched on one of the blocks, overlooks the zoo. This statue is unusual (it has a few flower-shaped blotches on its coat, not just stripes) and there’s some speculation that it was recycled from a life elsewhere where it played a very different role. Maybe it was previously a prehistoric cat in a theme park or a heraldic symbol or something; I’d love to know.

Caption: that’s a really interesting statue. Image: Darren Naish.

The zoo is not large and every exhibit can be observed and inspected within three hours or less. I visited, together with my son, Will, during February 2019, when the weather was less than ideal for zoo-going (it was cold, and foggy and raining).

Caption: the two white lions that were at the zoo when we visited in early 2019: Frosty (left) and Casper. Images: Darren Naish.

Among the most exotic and exciting of animals at the zoo are the white lions, two of which were present at the time of my visit (though read on): the male is called Casper, while the female is Frosty. White lions are recessive leucistic mutants and were first documented at Timbavati, South Africa, during the 1930s. They became better known to naturalists and zoologists at large thanks to Chris McBride’s book The White Lions of Timbavati (McBride 1977). The two I saw at the zoo are the only individuals I’ve ever seen. They’re not pure white, more like very pale blonde. Both came from West Midland Safari Park. Two other individuals – Vigo and Kumba – have joined the Isle of Wight Zoo since the time of our visit and both come from life in a travelling circus. Both were rescued by the AAP Animal Advocacy and Protection Primadomus, and you can read about the complex story of their rescue and transport to the zoo here. Both animals were castrated at an early age and this explains their unusual size and manelessness. I’ll have to revisit the zoo to see them some time, they sound fascinating.

Caption: Casper the white lion, exhibiting flehmen response, February 2020. Image: Darren Naish.

There are also a few smaller mammals at the zoo, namely Bennett’s wallabies Macropus bennetti, some extremely floofy Common raccoons Procyon lotor, South American coatis Nasua nasua (named Grant and Sattler, tell me you get the reference), Meerkats Suricata suricatta, a Crested porcupine Hystrix cristata, Lesser hedgehog tenrec Echinops telfairi, both African pygmy Atelerix albiventris and European Erinaceus europaeus hedgehog and several primates. The raccoon I watched (one of three) spent part of its time in its little house but eventually appeared for us and did the famous food-washing thing in textbook fashion. I’m sure that raccoons are tremendously familiar if you’re from certain parts of North America, but the same can’t be said of the UK, and indeed raccoons are not at all frequently encountered in British collections: I’ve only ever seen two or three, and I’m a regular zoo-goer.

Caption: one of the several raccoons at the Isle of Wight Zoo. So floofy. Image: Darren Naish.

A domestic animal section in the middle of zoo is home to especially big rabbits (they belong to a breed called the Giant European), goats, guinea-pigs, ferrets and pot-bellied pigs. Will and I were sufficiently impressed by the rabbits to spend some time looking at them. There’s supposed to be a single captive mouse in the zoo – called Mouse – who you can sometimes see rummaging in bags of hay, digging under turf or listening to the music of James Blunt. I see.

Caption: Harley (I think), a hybrid capuchin. These monkeys sometimes look like grumpy little people wearing thick fur coats. I’m following recent taxonomic proposals (Lynch Alfaro et al. 2012) in putting the ‘robust capuchins’ within Sapajus, and outside of Cebus. Image: Darren Naish.

The primates on show are Common marmosets Callithrix jacchus, spider monkeys, capuchins, Vervets Cercopithecus pygerythrus and members of six lemur species. The zoo’s spider monkeys are hybrids (between Black Ateles paniscus and Brown A. hybridus spider monkey) and the oldest of them are in their 30s and 40s. The zoo’s three capuchins were all born at the zoo and are also hybrids between Brown Sapajus apella and Black or Black-horned S. nigritus capuchin. The oldest of them – Timmy – is an impressive 31 years old. The hybrid nature of these various monkeys shows how different the philosophy on captive breeding was just 30-ish years ago.

Caption: spider monkeys at the zoo, February 2019. I’ve taken lots of photos of captive spider monkeys, usually because I’m trying to capture pictures where they look like de Loys ape. Images: Darren Naish.

The zoo also keeps a few birds, including owls and parrots, but the real highlight – the jewel is the crown – is the amazing…. Meller’s duck Anas melleri, a member of the mallard complex which I’ve written about on a few previous occasions. The zoo has five of these birds (all of which have very Madagascan names: Ambanja, Betafo, Itasy, Melaky and Sava) and is involved in the global conservation effort to replenish the population of this declining and endangered species. They successfully bred at the zoo in 2012.

Caption: could it be that I have before me…. a Meller’s duck? Such a fine bird. There’s a TetZoo review of knowledge on this species here at ver 2. Image: Darren Naish.

There’s also an education centre which also doubles as the home for various small animals kept in vivariums and cages. They include various insects, molluscs, amphibians and reptiles as well as a mouse lemur who only has one eye.

Caption: we’ve all seen Green iguanas Iguana iguana before (hey, the ‘Green iguana’ of tradition is a species complex that needs major revision). But they’re so great to look at that I never get bored of them. This big male was in the middle of eating. Image: Darren Naish.

While space is limited and the zoo is not large, you can see from some of the photos here that the larger animals have decent-sized enclosures. We were also told (while on a tour) that the cats are given a different form of behavioural enrichment every single day. On the day of our visit, the tigers had been given cardboard boxes and were carrying them around in their mouths. They seem to enjoy biting the cardboard and ripping the boxes to bits.

Caption: you can - in part - judge a zoo by its constructed installations. I approve of this rocky observation area for the tiger enclosures, opened by Chris Packham in 2004 (TV presenter Chris Packham is the partner of Charlotte Corney, the Wildheart Trust’s founder). Image: Darren Naish.

The Isle of Wight Zoo is not a spectacular or beautiful zoo. But it’s a good zoo, performing good work in terms of both animal rescue and welfare, in promoting connectedness between people and the natural world, and in conservation and education. It and its efforts should be supported, and I definitely recommend a visit if you’re interested in any of the animals I’ve mentioned here.

• Selection of species: 3 out of 10 (but see caveats)

• Zoo nerd highlights: white lions, hybrid capuchins and spider monkeys, raccoon, Meller’s duck

• Quality of signage: 7 out of 10

• Value for money: 9 out of 10

• Overall worthiness: 10 out of 10

My writing and research is dependent on crowd-funded support. Thanks to those whose patronage made this article, and the others you read here, possible. Please consider assisting me if you can, thank you!

For previous TetZoo articles on zoos and other relevant topics (I’m now routinely linking to the wayback machine versions, since those at SciAm have been ruined, or are paywalled. Yes, paywalled), see…

• ‘Revising’ the Siberian tiger, February 2009

• Lo, for I have seen the Meller's duck, and it was good, August 2009

• Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016

• Tet Zoo Reviews Zoos: Colchester Zoo, October 2017

• The Tiger Subspecies Revised, 2017, November 2017

Refs - -

Lynch Alfaro, J.W., Silva, J. S. & Rylands, A. B. 2012. How different are robust and gracile capuchin monkeys? An argument for the use of Sapajus and Cebus. American Journal of Primatology 74, 1-14.

McBride, C. 1977. The White Lions of Timbavati. Paddington Press, New York and London.

Tetrapod Zoology's 14th Year of Operation, 2019 in Review

Once again, it’s January 21st – hello, January 21st – which means that it’s Tetrapod Zoology’s birthday, or blogoversary or whathaveyou.

Caption: a montage of TetZoo-themed things relevant to 2019, read on…

TetZoo started life in 2006 and we saw last year how it became a teenager on its 2019 birthday. Today, TetZoo has hit the big 1-4. 14 years old. In keeping with tradition, let’s now look at the year’s TetZooniverous adventures, the caveat as always being that you should stop reading now if this sounds like it’s going to be too introspective. Because it will be. One final warning: this article is realllly long, I could have split into four or five different parts (but I didn’t want to).

Caption: we’ll start here. It depicts Joschua Knüppe, was taken at TetZooCon 2019, and was modified by Armin Reindl. I’ve lost track of who took the original, sorry.

From my own idiosyncratic perspective, TetZoo is – approximately speaking – as active as ever. I still manage to publish a few articles a month and report and discuss things relevant to my interests and thoughts, the site’s visibility and content continues to win me paying employment, its articles remain a (hopefully) valuable source of information on various arcane zoological topics, and – even in the age of Twitter and Instagram (I’m findable on both as @TetZoo) – it has a healthy community of regulars who help keep the site alive with discussion and comments.

Caption: TetZoo Towers, the scene of our story’s Act 1. Yikes, it was a mess at the time this photo was taken… Image: Darren Naish.

January 2019 started with me returning from a consultancy job in China – I work on occasion for Don Lessem’s DinoDon company – which I feel I’ve written about already. I visited Dinosaur Isle at Sandown (one of several trips made there during the year) for on-going work on Wealden theropod dinosaurs with Neil Gostling, Chris Barker and colleagues.

Caption: life-sized dinosaur models at Dinosaur Isle, Sandown. At left, one of Andrew Cocks’s Eotyrannus models (with cannon-fodder ornithischian in mouth). At right, the famous Dorling Kindersley Caudipteryx, with Luis Rey colour scheme. It was in a weird, crooked pose until Luis recently corrected it. Images: Darren Naish.

I also visited the Pulhamite Garden at Holly Hill. Pulhamite gardens – named for their designers, the Pulham brothers – are landscaped Victorian features, designed to include replica waterfalls, grottos, caves and so on. John Conway and I recorded a few episodes of the podcast and released episode 70… which we’d actually recorded back in September 2018. Yeah, we’ve had to abandon any plans to record episodes of the podcast at all regularly, not through choice but because workload no longer allows. And when we do get to record an episode, there’s no longer time or opportunity to edit it. Plus John is very, very lazy.

Caption: a cave - not a natural one, but one made by people - in the Pulhamite garden at Holly Hill, Hampshire, UK. I love this stuff. Reminiscent of the Prehistoric Court at Crystal Palace, of course. Image: Darren Naish.

The TetZoo review of 2018 (I mean, the 13th birthday article) was published at the end of January, a bit later than planned. Episode 71 of the podcast was released in early February and episode 72 – a Loch Ness Monster special – later in the month. A TetZoo article reviewing some recently-ish published books was published, and my recollections of the Dinosaurs Past and Present exhibition of the late 1980s and early 90s was published too. A technical paper I contributed to – on a Late Cretaceous eggshell assemblage from Romania, incorporating the eggs of several reptile species – was published in Scientific Reports in February (Fernández et al. 2019), and I wrote about it here at TetZoo. Will and I went to the Isle of Wight Zoo at the end of February. I mean to write about it. Internet potoos were also covered at TetZoo in February.

Caption: OH MY GOD IT’S MELLER’S DUCK. Yes, they have them at the Isle of Wight Zoo. Image: Darren Naish.

Also during February, I spent time at the BBC’s Natural History Unit – something that would take up an increasing part of my time for the rest of the year (no, I can’t talk about it) – and made tiny, incremental progress on the Eotyrannus monograph (it’s finished, and has been for months, ‘all’ I’m doing is making the post-review changes). My article on OroBOT appeared in BBC Focus magazine (Naish 2019a; an online version is here).

Caption: OroBOT! A mobile, articulated robot replica of Orobates. From the BBC Focus magazine article here.

Caption: the arrival of frogs and frogspawn in the early part of the year is always a significant annual event, and generally occurs round about the second week of February. Two clumps of spawn were produced in 2019; many of the tadpoles made it to froglet stage and some over-wintered and are still in the pond right now (January 2020). The species here is the European common frog Rana temporaria. Images: Darren Naish.

Books. The events of 2019 meant that I wasn’t able to write any new books during the year, nor finish any of the ones I’ve started. A few things happened though. The Dorling Kindersley book that I co-wrote with Chris Barker – part of their ‘What’s Where on Earth’, fully titled What’s Where on Earth: Dinosaurs and Other Prehistoric Life (Barker & Naish 2019) – appeared in print in March. It’s been very warmly received in reviews. I did an interview on the book (and dinosaurs in general) for First News newspaper.

Caption: new books for 2019. A new edition of Dorling Kindersley’s Dinosaurs & Prehistoric Life appeared during the year: I helped with the update but have yet to see a copy. What’s Where on Earth: Dinosaurs and Other Prehistoric Life (Barker & Naish 2019) also appeared.

The Japanese edition of Dinosaurs: How They Lived and Evolved (Naish & Barrett 2018) arrived. This version of the book makes me appreciate Japanese respect for detail and intricacy: the cover features several of the cladograms I produced for the book, the boards of the cover are decorated with a beautiful image of a moa skeleton, and there are additional details like an Erlikosaurus on the book’s spine. I drew squamates for the Big Book, a huge project that’s still ticking away in the background (cough cough).

Caption: cover views of the new Japanese edition of Dinosaurs: How They Lived and Evolved (Naish & Barrett 2018).

Moving to March, I worked on a TV series for Wall to Wall, commissioned by Netflix. I may or may not talk about it when it’s out. An article on alternative timeline dinosaur evolution – featuring interview comments and thoughts from me – appeared in the Italian Focus magazine (Camardo 2019).

Caption: opening spread of Giovanni Camardo’s 2019 article on alternative timeline dinosaur evolution. I’ve said it before but it’s worth repeating: James Kuether’s dinosaurs are everywhere these days.

The dinosaur models I worked on in China at the start of the year were delivered to their new home in New York’s Bronx Zoo and put on show. Articles published at TetZoo during this part of the year include those on the cautious climber hypothesis, and the first and second of my Nessie-themed book reviews. I attended a beach clean event in late March, and as usual picked up many kilos of discarded plastic crap otherwise contaminating the environment. I’ve been doing litter-picks at beaches for years now. Things haven’t improved but have steadily gotten worse. The end of much of the natural world is in sight.

Caption: imagine trying to clean a beach like this of its plastic pollution. It isn’t going to happen: no-one is ever going to remove all of this waste. Chessel Bay, Southampton, March 2019. Image: Darren Naish.

I went to another local zoo (Marwell) in mid-April, highlights of this trip being lemurs in a pile, a rhino gang, good views of Mountain zebra Equus zebra, Crocodile monitor Varanus salvadorii and Blesbok Damaliscus pygargus phillipsi.

Caption: Marwell Zoo has so many great animals. At left, Crocodile monitor Varanus salvadorii in the new tropical house. At right, Mountain zebra Equus zebra (note the dewlap). Images: Darren Naish.

TetZoo articles from this time included those on sleep behaviour in non-human animals, phyllostomid bats, cocks-of-the-rock, and my fondness for Usborne’s 1977 All About Monsters. My latest technical contribution to the cryptozoological literature appeared (Paxton & Naish 2019), this being an article in which Charles Paxton and I aimed to determine whether popular knowledge of Mesozoic marine reptiles might have influenced 19th and 20th century sea monster sightings. We concluded that they likely had, to a degree. This research was covered here at TetZoo; Mike McRae also wrote about it here at ScienceAlert. I spent time at one of my local patches, Telegraph Woods.

Caption: while at Portsmouth Comic Con in May, I got to see several of Roger Dicken’s movie models; here’s a sauropod, I think from The Land That Time Forgot. Images: Darren Naish.

May was busy. I visited the Royal Veterinary College for a secret project I can’t talk about and, outside of work, attended Portsmouth Comic Con and Exbury Gardens, went to the several famous geological sites of Durdle Dor and its surrounds on the Dorset coast (my parents spend part of the year there), and visited the new Dogstival fair – a massive, dog-themed event devoted to everything about dogs – in the New Forest.

Caption: some summertime shots of the New Forest, UK. It really is the most spectacular location and I love spending time there. Images: Darren Naish.

On the subject of dogs, late May was difficult…  I and my family said goodbye to Willow on May 20th, an event which inspired me to write about Willow and her life. I made the mistake of thinking that I’d be able to launch back into work and carry on as if nothing had happened; I should have taken the rest of the month, at least, off work. Other articles appearing at TetZoo during May include those on the creatures of Star Wars, palaeoartistic depictions of Styracosaurus, birdwatching in China, and cases where animals have died after they’ve been hit by falling rocks or trees.

Caption: I visited Bob Nicholls of paleocreations.com during summer 2019. Wow, he has some cool stuff. One of the psittacosaurs is for sale… Images: Darren Naish.

Hunting Monsters and the tabloid press. In early June I attended, and spoke at, the Cheltenham Science Festival. My talk was the Hunting Monsters one, initially prepared to promote the book of the same name (Naish 2017) back when it was new. The talk is on the history of cryptozoological theorising, on the reliability or otherwise of people as reporters of information, and on how the supposed targets of cryptozoology – ‘cryptids’ – might be explained. I signed and sold a bunch of books and had a great time, but an interesting thing happened after my talk.

Caption: books and book signing at the 2019 Cheltenham Science Festival. Jules Howard books make a guest appearance. Images: Darren Naish.

Two journalists – one from The Sun, one from The Daily Mail – were in the audience, and both asked if I thought that the lack of compelling photographic data (especially in an age where huge numbers of people now carry cameras with them most of the time) was essentially the death-knell for the existence of monsters. While things are not necessarily as simple as all that (not all animals can be photographed given the brief duration of many encounters, camera-phone photos are not necessarily good enough to be convincing to those with critical faculties, and so on), I do think that the rarity if not absence of photographic data for most cryptids counts for something, and I hence agreed with this contention. Both journalists were interested, quizzed me further on this point during the book-signing event, and said that this point would be used as the ‘hook’ in the pieces they were going to write. Remember: this idea was raised, de novo, by them and wasn’t mentioned or alluded to in my talk.

Caption: quality, genius journalism from the rag known as The Sun, oh what fun.

How, then, did the journalists concerned deal with this information? Well, you can see for yourselves from the screengrabs used here: the whole thing was spun in the weirdest direction, the take being that I’d only just concluded that cryptids aren’t real, and that I’d hence admitted to wasting about 20 years of research (as if monster-hunting is all I’ve done across the better part of my adult life). Needless to say, this is a nonsense interpretation of everything I’ve done and published. I wrote a response (initially as a series of threaded tweets); you can read it here.

Things kicked off from here, by which I mean that there was substantial further media interest. I turned down most requests for interviews (I’m at that point in my career and experience where I no longer see any benefit in making unpaid appearances on radio or TV. The claim that media exposure helps you, or is useful, is bullshit in my line of work) but did – under duress – do a brief thing for Channel 5 news (you can watch it here, if you want). So I learnt a lot from this, the main thing being that I’d underappreciated how vile and disgustingly biased and manipulative British tabloid journalism mostly is. Newspapers like The Sun approach issues with a rancid anti-intellectual agenda, with a clear intention to inspire and promote disdain, hate and ill will and – while an article written to make me look like an idiot has no national or international relevance or importance and won’t be remembered by anyone excepting myself – the impact that these redtop rags have on important issues (like Europe’s refugee crisis and the infinitely bad Brexit shitstorm) can’t be understated. Everyone knows this already, of course; I merely wanted to state it in my own words.

Caption: at the 2019 Dinosaurs and Art event - hosted at Oxford University Museum of Natural History - David Button uses the mounted Iguanodon bernissartensis replica to make a point about iguanodontian anatomy and posture. Image: Darren Naish.

Whale Watching, Fortean Times, Monsters of the Deep, Anglesey. Also in June, I watched Godzilla: King of the Monsters and was sufficiently inspired to write about it. I attended Will Tattersdill’s Dinosaurs and Art event at the University of Oxford and enjoyed talks by David Button and Verity Burke. I visited Bob Nicholl’s studio in South Gloucestershire (see montage above) and got to see a great deal of amazing in-prep, embargoed work, what an honour. My reminiscences of Lyall Watson’s Whales of the World, some thoughts on books about woodpeckers and my thoughts on Mark Witton’s The Palaeoartist’s Handbook were published at TetZoo.

Caption: you need a really good camera to take good photos of distant whales, and I don’t have one. So what I’d sometimes do, as a temporary record of recording what we’d seen, was photograph other people’s camera screens. Here, you can see someone else’s photo of two Cuvier’s beaked whales. Image: Darren Naish.

During July I went on a whale-watching trip in the Bay of Biscay. This was a fantastic experience (bar the desperate rush to get to the ship on time, I only just made it) where we saw hundreds of dolphins, several Fin whales, Minke whales, Cuvier’s beaked whales, Harbour porpoises and so on and I fully intend to go again as soon as opportunity allows. I wrote about the trip here; thanks again to Alex Srdic for use of his photos.

Caption: a scene from the Pont Aven, July 2019. Our progress across the Bay of Biscay is charted, and we keep tally of the cetaceans (and other interesting animals) we see. Image: Darren Naish.

Topics covered at TetZoo included European cave art, British journalism’s misplaced and weird hatred of gulls (see above re the British tabloid press) and dunnocks, and I also published an article reviewing the FIRST YEAR of TetZoo at its home here at tetzoo.com. I drew lots of passerines and snakes for my textbook and TetZooCon tickets went on sale.

Caption: here’s something that’s kind of a big deal. I worked - as consultant - on the second edition of Don Lessem’s Ultimate Dinopedia book, published 2017. And - look - here it is, making a guest appearance in season 3 of The Handmaid’s Tale (in an episode that I watched in July 2019). In the episode, Emily reads part of the section on Spinosaurus (but changes the wording slightly relative to what’s actually in the book).

A really interesting TetZoo-relevant article appeared in Fortean Times in July: namely, a whole article on Hunting Monsters (Naish 2017), published as part of their ‘building a Fortean library’ series. The article was penned by The Hierophant’s Apprentice (there’s a long backstory there which’ll mean something to followers of Forteanism) and emphasises the book’s value as a sceptical review of cryptozoological claims and ideas (The Hierophant’s Apprentice 2019).

Caption: Hunting Monsters is written about in Fortean Times 382, what an honour.

On to August. I don’t know if I’ve mentioned here my in-prep museum exhibition – Monsters of the Deep – which will open in March 2020 at the National Maritime Museum in Falmouth, Cornwall. At the time of writing it’s near-finished and we’re about ready to go, but in August there was lots to do and I spent time in Falmouth, pulling things together. More on the exhibition below.

Caption: Red-billed choughs Pyrrhocorax pyrrhocorax on Anglesey. As usual, my photos aren’t the best, but at least they show what I saw. Choughs are among the oddest of corvids. Images: Darren Naish.

I travelled from Falmouth to Wales, meeting en route with my family, for a brief holiday in Anglesey. We had a fantastic time and watched Common bottlenose dolphins Tursiops truncatus and Grey seals Halichoerus grypus (both within just a few tens of metres of our residence), Red-billed choughs Pyrrhocorax pyrrhocorax, a huge raft of Goosander Mergus merganser and a ton of seabirds.

Caption: a Goosander raft in the Menai Straits, Anglesey. Image: Darren Naish.

At Breakwater Country Park we visited the outdoor Charles Tunnicliffe gallery, which was a great thrill to me as he’s one of my favourite natural history artists, the works he contributed to (Ladybird Books and PG Tips teacards) being among my earliest influences. The Anglesey Sea Zoo was also great; I used the life-sized leatherback model in an effort to make a homage to the 1988 photos of the giant Harlech leatherback, a century-old specimen that stranded (after drowning in a fishing net) in Gwynedd, Wales, in September 1988 and is today on show at the National Museum, Cardiff. We also visited Ironbridge in Shropshire where I managed to pick up a (super cheap!) copy of J. A. Moy-Thomas’s 1939 book Palaeozoic Fishes.

Caption: my best efforts to mimic what happened in photos depicting the famous Harlech Beach leatherback. This model is on show at Anglesey’s Sea Zoo.

My article on British tabloid journalism’s approach to KILLER SEAGULLS appeared in BBC Science Focus magazine (Naish 2019b) and a reprinted article on the rise of dinosaurs in the Triassic also saw print (Naish 2019c). TetZoo articles of August included those promoting TetZooCon 2019, my review of Angus Dinsdale’s Loch Ness Monster book on his father (Tim Dinsdale), and a report on whale watching in the Bay of Biscay.

Caption: I took so many photos on Anglesey that I have too many to share, but here’s one more. Green spaces are of vital importance to our well-being, we must do everything to preserve them. The humans are Emma, Will and Toni. Image: Darren Naish.

King of the Sex Lakes. While out shopping I discovered that Brian Ford’s wretched book Too Big to Walk had succeeding in winning a second edition. Here’s the explanation for the ‘dInOsAuRs NeEdED sEx LaKeS’ story – wherein it was, apparently seriously, proposed that dinosaurs needed lakes for the purposes of mating, and that a drying up of said lakes in the Late Cretaceous caused dinosaur extinction – which hit the newswires earlier in the year (in May): it was evidently released to drum up interest in Ford’s alt-universe dinosaur project. Ford is a science writer who seems to regard himself as the ultimate expert on everything and, prior to his efforts to reform our understanding of dinosaurs (he thinks that all non-bird dinosaurs were aquatic), might be best known for his takes on spontaneous human combustion and alternative energy weapons.

Caption: a now iconic image. Nothing says science like a full run of Encyclopedia Brittanica.

I’ve already written too much about him (Naish 2012, plus this 2015 article at TetZoo), and at least some of you will know that I gave a talk at his own book launch (held at Conway Hall during May 2018). I did my best in that talk to counter his writings on dinosaurs and out them as the steaming pile they are. Anyway, said second edition of Too Big to Walk includes various updates relative to the first edition, the most interesting of which are those pertaining to the events of Conway Hall. Ford’s take on what happened is dishonest and inaccurate and I did my best to more accurately report events in a series of threaded tweets which you can read here.

Caption: Mr Ford talks about Riley Black on a cruise liner (from this 2015 TetZoo article). This is an old photo that comes from TetZoo ver 3: I try not to link directly to the SciAm host of ver 3 but to the wayback machine versions of the respective articles, firstly because SciAm have removed most of the images; secondly because they’ve paywalled them too. Image: Darren Naish.

Too Big to Walk, incidentally, currently scores 2.5 on Amazon and the only positive reviews appear to have been written by Mr Ford’s friends and allies. This is despite the fact that Ford (and/or his publishers) successfully lobbied to get several fair – but harshly negative – reviews written by qualified palaeontologists removed for ‘being biased’. As I’ve said before, I don’t want to write or talk about Brian Ford and his aquatic dinosaur nonsense again – what a waste of my time and effort – but I’ll continue to do so if I have to.

Caption: the ‘sex lakes’ idea was mocked hilariously throughout the year. I am nothing to do with these images, widely shared online.

It is especially amusing to see Mr Ford lie about the way his claims have been received within the palaeontological community. He’s claimed (on one of his facebook groups) that young palaeontologists “love” his work, that his stuff is only being resisted by the ‘old guard’ – who are busy squatting on their piles of accrued palaeo-dollars at the tops of their ivory towers – and that scientists far and wide are accepting his writings with open arms. When challenged, he can’t name any such scientist, however. I asked on Twitter if anyone out there really “loves” his stuff. The responses were pretty amusing, in that the main “love” for Ford’s view of dinosaurs comes from its potential as meme-worthy nonsense that’s fun to mock.

On that note, early 2020 saw the release of Lemme Splash!, a game in which your aim is to get two amorous, love-struck sauropods to the sex lakes. An excellent review, with some exposition, is provided here at Dino Dad Reviews. “Brian Ford, this is your legacy”.

Conference World. September and October are conference season, and September kicked off with PopPalaeo (properly: Popularising Palaeontology), hosted at King’s College, London, and arranged by Chris Manias. The entire meeting was devoted to palaeontological representation via media, and my own talk was on Dinosaurs in the Wild. It’s online here. Ilja Nieuwland brought along one of the recently discovered replicas of the original Hydrarchos skulls and there was a public engagement event involving an art exhibition and some public talks.

Caption: a Dinosaurs in the Wild Tyrannosaurus model observes events at PopPalaeo 2019. Image: Darren Naish.

The season’s second conference was the 67th SVPCA, this year held at Ryde, Isle of Wight. Because I only had clearance to go at the very last minute, I didn’t have time to throw a talk together. But I really wish I had, since my continuing (and seemingly never finished) research on Wealden theropods would have been absolutely relevant given the biases of the audience. The meeting was made memorable by my locking our entire party out of our shared accommodation, what fun.

Caption: some scenes from the 67th SVPCA, Isle of Wight. At left, a 3D-printed Neovenator skeleton. At right, a table-load of John Sibbick originals. Yes, ORIGINALS. Images: Darren Naish.

The big TetZoo-relevant event of the year is TetZooCon of course, and September is the month where things really have to be sorted out (John’s hectic schedule demands that we leave everything to the last minute). And thus it was that we discovered that The Venue – our, err, venue – had double-booked the room we needed, their suggested solution being that we hold the event at one venue on the first day but at a second venue on the second. Needless to say, this was unworkable and another solution would be needed. We got things resolved eventually, but what a mess. Even after five years of running these events we have yet to find a venue which is (a) affordable and (b) actually interested and reliable when it comes to organisation and communication.

Back at TetZoo, I published my review of Phil Senter’s Fire-Breathing Dinosaurs? as well as parts 1 and 2 in the extreme cetaceans series. Episode 74 of the podcast was released.

Caption: it turns out that Brisbane is absolutely full of these amazing animals. I photographed every individual I saw. They were Australian water dragons Intellagama lesueurii. Image: Darren Naish.

During early October, my wife Toni and I flew to Scotland for the wedding of our friends Jeff and Femke (I’m fundamentally opposed to the concept of short-haul flights but… ugh… if you need to get to Scotland from southern England, flying is massively cheaper than going by train, ffs). I was only back a few days before I attended conference number three: the 79th Society of Vertebrate Paleontology meeting, this year hosted in Brisbane, Australia. This was my very first trip to Australasia. I saw loads of great animals (my article on the birds I saw – published in mid November – is here), hung out with friends old and new, and obtained books and animal figures that aren’t ordinarily available back home in the UK. I attended the meeting for the talks and posters on Cretaceous dinosaurs, pterosaurs and marine reptiles but it was made especially memorable by the many excellent presentations on Cenozoic marsupials, lizards, crocodylians and birds. A dinosaur-themed presentation that inspired a lot of discussion was Kayleigh Wiersma and Martin Sander’s on the claimed presence of beak-like tissues in sauropods; I was one of the several palaeontologists interviewed by John Pickrell for his Science article on the proposal.

Caption: scenes from the excellent Queensland Museum, Brisbane. At left, the famous Perentie Varanus giganteus specimen that died while swallowing an echidna (there’s an old TetZoo article about it here). At right, a real live wombat! Images: Darren Naish.

Finally on the conferences, conference number four – TetZooCon 2019 – happened on the weekend of 19th and 20th October. It was the biggest and best TetZooCon so far and included a dinosaur palaeobiology session and roundtable, a natural history film-making panel and section of talks, Mike Dickison’s Wikipedia workshop, several book signings, a dedicated palaeoart event and exhibition, and a proper merchandise area. The TetZoo write-up is here. Other reviews of the meeting can be found here at LITC (and here’s part 2), here at Luis Rey’s blog, here at Talita Bateman’s blog, here at Steve Allain’s herpetological blog, and here at Albert Chen’s Raptormanics.

Caption: at left: Lauren, Laura, Sam and Kannan at London’s Zoo Reptile House. At right: your humble author with a giant penguin. Always with the giant penguins. This one might be a little too giant. Images: Darren Naish.

A bunch of us went on a post-TetZooCon fieldtrip to ZSL London Zoo on the Monday after the convention. We saw nearly all of the animals. At the time of writing, John and I are in the earliest stages of discussion about TetZooCon 2020. We still don’t have the finances resolved from last year, so I can’t yet be sure that it’ll definitely be happening. But it probably will.

Caption: at left: Dr Cox/BiPolarCosplay as SkekSo, Skeksis Emperor. At right: Naish and Reddish, together at last.

I got to see the first episode of the new BBC natural history series Seven Worlds, One Planet in the cinema and attended London ComicCon with the kids. Highlights included meeting Reddish from Don’t Hug Me I’m Scared, Spawn and Violator, a Skeksis, a wookiee and a Hoth wampa. An entire episode of the Natural History Channel Podcast devoted to TetZooCon 2019 was released in November 2019 and can be found here. At TetZoo, my review of Robert France’s book on the New England Sea Serpent was published.

Caption: some Naishes (Will at left) with a Wampa; junior Hoth rebel at far right.

The freshly renamed pterosaur Targaryendraco was published at the end of November; I work on pterosaurs sometimes (I certainly have enough unfinished manuscripts on them kicking around) and was quoted in this Nat Geo article by John Pickrell. I actually told John a bunch of stuff that (for obvious and understandable reasons) didn’t make it into the final article, namely that Targaryendraco was being worked on by a second team who – arguably – could be said to have ‘academic priority’ on this project, and that the systematics and taxonomy of ‘ornithocheiroid-type’ pterosaurs has been slowed and stalled for decades by the efforts (or otherwise) of a certain pterosaur worker who shall remain nameless. FINAL EVIDENCE that the Phylonyms: A Companion to the PhyloCode volume will appear in 2020 – only a few decades too late – was received in November. I have one contribution in said volume. I should have three but – funny story – two of them were rejected after having been accepted, substantially revised in a loooong running series of exchanges, and listed as ‘in press’.

Caption: my time in Bristol means that I’ve made numerous visits to the Bristol Museum and Art Gallery, which has a ton of excellent stuff. Here’s a dodo model, and an Archaeopteryx. Images: Darren Naish.

I went on a Climate Strike march in Bristol… I still can’t decide whether these events are a total waste of time or not: when it comes to the action that’s so urgently required, we’re still being dismally and catastrophically failed by the so-called leaders of our various countries.

Caption: a Climate Strike march in Bristol, November 2019. We want our governments to instigate change. They mostly want to carry on as normal, because who cares about the future. Image: Darren Naish.

A series of TetZoo articles on ‘alternative timeline dinosaurs’ appeared (part 1, part 2, part 3), which was good (and not entirely coincidental) timing since a BBC World Service radio show on the very same subject was released in November, and featured myself in addition to Memo Kösemen, Anjali Goswami, Elsa Panciroli and Nicola Clayton. I got to see one of my favourite bands – Metronomy – live at Bristol’s O2 Academy. And, with the family, I visited Birdworld in Surrey, home to one of the UK’s most exciting bird collections.

Caption: an ‘alternative timeline smart dinosaurs’ montage, featuring various creatures written about in the TetZoo series. Images: Jim Limwood, CC BY 2.0 (original here), Norman (1991), John Sibbick/Norman (1985), Darren Naish, C. M. Kösemen, John McLoughlin, Matt Collins, Mette Aumala.

Caption: a selection of birds on show at Birdworld, Surrey. They have a lot of great stuff. Images: Darren Naish.

A spider story. While out one December evening at a busy restaurant, I saw a large dark spider on the floor, and – as you do – went to pick it up so that I could relocate it in a place of safety. Unlike any other spider I’ve previously handled, it immediately bit me hard, latching its giant fangs into my left thumb and leaving two small bleeding holes in my skin. I’d been envenomated, and the wound hurt for the next few hours. The pain was about similar to that of a bee sting. The spider was Segestria florentina (the Tube web or Mouse spider), a species I know well but have never previously been bitten by. Segestria, incidentally, isn’t native to the UK but is now so widespread that it’s an established part of our fauna. I captured the spider in a small bucket and released it outside. Remember that animals which bite you are (ordinarily) terrified and in fear of their lives, not ‘angry’ as is so often stated.

Caption: Flame the dragon, on this occasion being most upset about the look of the sky. Image: Darren Naish.

It was also in December that we went live on the advertising for the Monsters of the Deep exhibition which I mentioned earlier. This has been a big part of my life for the last few years and rest assured that I’ll be writing about it once the exhibition is officially open. It’s mostly about European sea monster lore and covers medieval ideas about the creatures of the sea, mermaids, kraken, the scientific exploration of the oceans and cryptozoology. There isn’t currently much online content about the exhibition but, right now, you can get a basic idea of what’s going on here.

Caption: Monsters of the Deep teaser - more info is available here, and watch the National Maritime Museum, Falmouth Twitter account too.

Pressures of time mean that nothing has been published at TetZoo this month – as in, January – bar this article and the one on the Loveland Frog, the latter written as a tie-in to our ongoing efforts to get Cryptozoologicon Volume 2 finished (the ‘our’ referring to me, John Conway and C. M. Kösemen). And that, just about, brings us up to date…

In keeping with tradition, it’s at this point that I list the TetZoo articles of the year by category, analyse said coverage with taxonomic bias in mind (how fair, or unfair, have I been to the various tetrapod groups?), and then self-flagellate given that the bias usually reveals massive skew to charismatic megafauna. Here we go…

Miscellaneous Musings

• TetZoo Bookshelf, February 2019, Part 1

• Sleep Behaviour and Sleep Postures in Non-Human Animals

• On May the 4th, Some Star Wars Musings

• Death by Falling Rocks and Trees

• Godzilla: King of the Monsters, Some Trivial But Monstrous Thoughts

• Mark Witton’s The Palaeoartist’s Handbook

• Five Famous Palaeolithic Rock Art Enigmas

• The First Year of Tetrapod Zoology Ver 4

• Announcing TetZooCon 2019 – the Biggest Yet

• Final Call For TetZooCon 2019

• The Sixth TetZooCon

Mammals

• The Cautious Climber Hypothesis

• The New World Leaf-Nosed Bat Radiation

• Lessons From a Dog: a Life With Willow

• Remembering Lyall Watson’s Whales of the World

• Whale Watching in the Bay of Biscay

• Extreme Cetaceans, Part 1

• Extreme Cetaceans, Part 2

• Extreme Cetaceans, Part 3

Non-avialan dinosaurs

• Recollections of Dinosaurs Past and Present, the 1980s Exhibition

• The Fate of Burian’s Styracosaurus

• Philip J. Senter’s Fire-Breathing Dinosaurs?, the TetZoo Review

Birds

• A Multi-Species Nesting Assemblage in the Late Cretaceous of Europe

• Potoos As An Internet Phenomenon

• Cocks-of-the-Rock, Extreme Cotingas

• Birdwatching in Suburban China

• The World’s Best Books on Woodpeckers

• Tell Me Something Interesting About Dunnocks

• Hacks Vs Wildlife: the Eternal Vilification of Gulls

• Suburban Birdwatching in Queensland, Australia

Cryptozoology

• Books on the Loch Ness Monster 1: Ronald Binns’s The Loch Ness Mystery Reloaded

• Books on the Loch Ness Monster 2: Gareth Williams’s A Monstrous Commotion

• Usborne’s All About Monsters

• Sea Monster Sightings and the ‘Plesiosaur Effect’

• Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness

• A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent

• Lore of the Loveland Frog

SpecBio

• Alternative Timeline Dinosaurs, the View From 2019 (Part 1)

• Alternative Timeline Dinosaurs, the View From 2019 (Part 2)

• Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums

Why Are Things Like This. Ok, there’s a bit to unpack here. Long-time readers will know that my aim at TetZoo is to achieve some sort of taxonomic balance whereby I write fairly and equally about the major tetrapod groups. Somehow I stupidly expect that this will magically just happen, which I know is ridiculously flawed and dumb, but there it is. Have I ever achieved said balance? There was a time when I felt I was getting close to achieving it but… no, no I haven’t. And as we can see from TetZoo’s subject coverage across 2019, I’m further away from it than ever, utterly failing across the year to cover amphibians, stem-mammals, squamates and so many other groups. This failure is mostly down to two things, and those things are things that make the ‘fair coverage’ failure somewhat ironic, and hopefully understandable. Forgivable, if you will.

Caption: for levity, some toys. Images: Darren Naish.

First thing: now that TetZoo is hosted at its very own site, I feel freer, more able to write about whatever relevant topic I like, however sensational or semi-serious. And I won’t lie: writing about the Loveland frog, alternative timeline dinosaurs or the Loch Ness Monster is easier for me than is writing about obscure skinks of New Guinea or turtle phylogeny. Those more technical, scientific topics – while worthy and of great interest to me – require so much more research and checking of the literature, and thus time. Time is the great constraint on everything I do. Oh, and money, but I’m not here to complain about that.

Caption: which of these two articles do you think brought in more visitors: the one about bigfoot’s dick, or the one about mystery skinks from Tonga? The answer may depress you, but may also be very predictable.

Second thing: tetzoo.com is still a relatively new site where I’m trying to build an audience. Both ScienceBlogs and SciAm – the two previous hosting sites for TetZoo – came with captive audiences of many thousands of readers, any number of whom would hop over to TetZoo when the interesting title of a TetZoo article appeared in a sidebar. This new hosting site doesn’t have that ‘captive audience’: I’m relying either on people who come here on a regular basis because they’ve been rewarded in the past (hello), or those who find the site while googling ‘what do we know about bigfoot’s genitals’, or whatever (hello to you too, please visit again). Ergo, part of my thinking has been that I should concentrate on some of the rather more sensational things I write about: experience and data shows that they do bring in more readers. If you doubt that, here’s a visitor graph showing what happens when I write about ‘humanoid dinosaurs of an alternative timeline’. People are fickle, and I’m trying to take advantage of that.

Caption: this is what happens when I blog about speculative zoology… I mean the peak at far right. So don’t judge me.

In fairness to myself, I should also add that my assorted dealings, projects and paying gigs over 2019 mostly revolved around cryptozoology, dinosaurs, speculative biology and sensational, weird animals, and not the more academic side of herpetology or vertebrate palaeontology. So there’s also some genuine justification for the biases in coverage we’re seeing here.

Caption: Andrew Dutt produced this masterpiece. It reflects John Conway’s great love of movies like Pacific Rim and the unstoppable force that is the Category V kaiju Petersquama. Image: Andrew Dutt, used with permission.

And that about wraps things up. This article is huge: about five times longer than a typical TetZoo article, and I did toy with the idea of splitting it up. I opted not to because I prefer to avoid dragging the birthday thing out; I’d rather move on to other things. Despite an ever-increasing workload and a continuing decrease in the time available to me for blogging (I’m writing this while sat on a train: I’ve just seen a group of Roe deer in a frozen field), my aim is to continue to publish new content as and when I can throughout 2020 and beyond. I thank you for visiting and reading TetZoo and hope that you’ll continue to do so. And happy 14th birthday, little blog.

Caption: another personally relevant thing I haven’t discussed here at all: 2019’s maturation of pond 2. It and its surrounds proved a total wildlife haven. I dearly hope this small green area persists after we move away…

For previous Tet Zoo birthday articles, see...

• Happy first birthday Tetrapod Zoology (part I)

• Happy first birthday Tetrapod Zoology (part II)

• Happy second birthday Tetrapod Zoology (part I)

• Tetrapods of 2007 (happy birthday Tet Zoo part II)

• Happy THIRD birthday Tet Zoo

• Tet Zoo = 4 years old today

• 2009, a year of Tet Zooery

• Four years of Tet Zoo: to infinity... and beyond!

• It is with some dismay that I announce Tet Zoo's first hemi-decade

• Tet Zoo 5th birthday extravaganza, part II

• Happy Birthday Tetrapod Zoology: SIX YEARS of blogging

• Happy 6th Birthday, Tetrapod Zoology (part II)

• Tetrapod Zoology enters its 8th year of operation

• Today marks NINE YEARS of Tetrapod Zoology

• Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review

• Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed

• Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015

• Today Is Tet Zoo's 11th Birthday

• The 12th Year of Tet Zoo

• The Tet Zoo 12th-Birthday Event, Part 2

• The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event

• Tetrapod Zoology Is A Teenager Now, January 2019

Refs - -

Barker, C. & Naish, D. 2019. What’s Where on Earth: Dinosaurs and Other Prehistoric Life. Dorling Kindersley, London.

Camardo, G. 2019. Se non si fossero estinti, avremmo vinto noi o loro? Focus 317, 51-55.

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.

Naish, D. 2012. Palaeontology bites back… Laboratory News May 2012, 31-32.

Naish, D. 2017. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Naish, D. 2019a. The walk of prehistoric life. BBC Science Focus 334, 52-55, 58-59 (online version here).

Naish, D. 2019b. Seagulls: are they getting more aggressive? BBC Science Focus 340, 38-39.

Naish, D. 2019c. How dinosaurs conquered the world. BBC Science Focus Magazine Collection 13, 42-47.

Naish, D. & Barrett, P. M. 2018. Dinosaurs: How They Lived and Evolved. The Natural History Museum, London.

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

The Hierophant’s Apprentice. 2019. Building a Fortean library No 47. There ain’t so such animal. Fortean Times 382, 56-57.

Lore of the Loveland Frog

At least some of you reading this will know my 2013 book The Cryptozoologicon, produced in collaboration with John Conway and C. M. Kösemen (Conway et al. 2013).

Caption: at left, cover of the 2013 book The Cryptozoologicon. At right, a scene from the book’s interior, depicting yetis in a Himalayan scene; by John Conway. Image: Conway et al. (2013).

The Cryptozoologicon is devoted to cryptids; that is, to mystery animals. After describing a given cryptid and proposing how it might actually be explained (sadly, most cryptids now seem to be sociocultural phenomena or the products of fakery or human error, not valid biological entities; Naish 2017), we go on to indulge in a bit of speculative zoology: that is, a bit of ‘what if’ speculation whereby we imagine ourselves inhabiting a parallel universe where cryptids are real (Conway et al. 2013)…

TetZoo regulars will know that a sequel to The Cryptozoologicon – it’s working title is The Cryptozoologicon Volume 2, duh – has been planned for some time, and we still aim to complete it ‘soon’. Which creatures will be covered in this soon-to-be-published work? I’m not saying, but the article you’re reading now concerns one, just one, of the cryptids we’ve included.

Caption: I really think that some artists have made the Loveland Frog look substantially too frog-like, and this is one of the most extreme examples. But, hey, it’s a nice and technically very competent piece of art. Nice to see Pioneer Dork being used as a scale bar. Image: artist unattributed, Cryptid Wiki (source).

Books on ‘mystery animals’ – cryptids and the like – include a veritable panoply of the weird, unbelievable and ridiculous. Among these, one of my favourites is the Loveland Frog (sometimes called the Loveland Lizard, but that’s just silly): a bipedal, vaguely reptilian animal, similar in size to a child, and supposedly encountered several times in the vicinity of Loveland, Ohio, USA between 1955 and 1972… and there are a few more recent claimed sightings too. The sightings describe greyish, bipedal creatures, said to have a frog-like head, bulging eyes, a leathery, reptile- or amphibian-type skin, and a standing height of between 3 and 4 feet.

The Loveland Frog has been discussed and revisited several times in the mystery animal literature, the tales recounted here having previously been published in Bord & Bord (1989), Newton (2005) and Shuker (2008)*. I especially like Bord & Bord’s (1989) section on the case because it includes the wonderful illustration you see below, produced by Ron Schaffner, and evidently based on a pencil drawing produced by one of the witnesses.

* Confession: I didn’t have Loren Coleman’s Mysterious America to hand while writing, nor W. Haden Blackman’s The Field Guide to North American Monsters. I understand that these works also include coverage of this creature.

Caption: Ron Schaffner’s evocative illustration of the Loveland Frog, very obviously based on the pencil sketch shown below. Note the suggestion of cranial horns, the rows of parasagittal spines and the oval eyes. Image: Ron Schaffner, from Bord & Bord (1989).

Sources that discuss the Loveland Frog most usually recount the observation of police officer Ray Shockey (though spelt Shocke in some sources) who stopped to observe a creature seen crossing the road at 1am on March 3rd 1972, when it was cold enough for the road to be icy. Shockey’s creature was in a crouching position but then stood erect and stared in Shockey’s direction before climbing the guardrail separating the road from the ground that slopes down to the Little Miami River (Newton 2005, Haupt 2015). Other officers later came out to check Shockey’s observations. There’s some talk of them finding scratch marks on the guardrail but efforts to locate photos verifying the presence of said scratches haven’t been successful (Haupt 2015).

Caption: A pencil drawing of the Loveland Frog, I assume that made by Ray Shockey (though I’ve been unable to confirm this; I found it, unattributed, at various sites online and haven’t seen it in print). The artist evidently had quite some skill.

An account similar to Shockey’s was made by another police officer – Mark Matthews – two weeks later, and this again involved the animal being encountered on the road at night and seen from a patrol car. Matthews was concerned as the creature stood up from a crouched stance and fired his gun at it – yee-haw!! ‘Murica!! – and seemingly injured the animal. Again, it climbed out of sight over the guardrail.

Caption: another depiction of Shockey’s frogish encounter, this time showing the creature with a sumptuous butt and disturbingly human-like physique. This image is widely available online but I’ve been unable to find the artist’s name.

But Matthews later claimed that none of this was accurate, that he’d actually seen a big lizard (an escaped pet iguana?), and that he’d augmented the story as a way of making his colleague (Shockey) seem like less of a nut. I really like the deep investigation of this account provided by Ryan Haupt for the Skeptoid Podcast (here; Haupt 2015). It provides lots of additional information and is clearly substantially more reliable than the recountings of events provided in standard cryptozoology- and paranormal-themed websites and publications. Haupt states that there’s what appears to be an email confession from Matthews whereby the account was dismissed as being ‘blown out of proportion’, though its authenticity (it – the ‘confession’ – appears to have originated from this 2001 article from X-Project Paranormal Magazine) is doubtful. An implication that a big lizard might have been seen must also be considered doubtful in view of the icy conditions (and cold temperatures) of the time. A big lizard would be hiding away somewhere, not out and about. Apparently, there’s a sketch that accompanies either Shockey’s or Matthews’s account (Newton 2005; though his text seems to combine both accounts into a single sighting); I assume it’s the pencil one shared above. A local farmer is also said to have seen a Loveland Frog shortly afterwards but details are hazy.

Caption: images showing all three of Hunnicut’s Loveland Frogs together are rare, but at least there’s this fine piece of work by John Meszaros. Image: Cryptids State-by-State, John Meszaros (source).

The oldest Loveland Frog story pre-dates these 70s one and concerns a sighting made on an Ohio roadside during the early morning of May 25th 1955. The story goes that a businessman or travelling salesman – sometimes said to be unknown and sometimes specifically identified as Mr Robert Hunnicut (Newton 2005) – was driving home from work when, at 3.30am, he saw three bipedal, greyish reptilian creatures, each about 3 feet tall. The witness stopped and observed them for a few minutes. In some versions of the story, the creatures were seen ‘conversing’, in some they were observed under a bridge, and in some one of them held a cylindrical or wand-like device above its head. This released sparks and was frightening enough that it inspired the witness to flee.

Post-1972, the Loveland Frog has been rare. There’s a 2016 event in which two teenagers – out playing Pokémon Go, apparently (this isn’t a sexual euphemism) – supposedly saw and even photographed what appears to be the creature. But as you can see for yourself, this event is almost definitely a hoax.

Caption: a still from the 2016 footage taken by Sam Jacobs and his girlfriend. The actual footage is exceedingly dark and this image has been brightened as much as possible (by the people at Fox19 News). Some think that the photo actually shows a lawn decoration with added lightbulbs. Image: Fox19 News (source).

Explaining the Loveland Frog, or trying to. Given the several peculiarities of the 1955 ‘three creatures’ account, it’s not surprising that some authors have sought to identify the Loveland Frog as an alien rather than an unknown denizen of Planet Earth. Assuming that Loveland Frog accounts represent actual observations, an explanation mooted by some authors is that they could be confused descriptions of escaped pet iguanas or monitor lizards (Bord & Bord 1989). This is hard to accept given the bipedal postures that witnesses reported, plus the descriptions don’t recall big lizards at all. At a stretch we might consider that big lizards in fleeting bipedal or erect-standing poses were witnessed, with substantial embellishment and confusion resulting in substantially modified descriptive accounts, but the cold temperatures present during some of the sightings also count against this idea.

Perhaps, some might suppose, these fleeting glimpses of big escaped lizards were inadvertently (or deliberately) combined in the minds of the witnesses with their prior knowledge about the ‘big frog monsters’ already said to inhabit the Miami River region. Such stories go back to the 1950s at least and it should be noted that a similar-sounding entity, the ‘Lizardman’, was reported during the 1970s from South Carolina, New Jersey and Kentucky. In other words, there seems to be lore in the region about such creatures… which might mean that any fleetingly-glimpsed, unidentified weird animal could morph into a monster of this sort in the memories of witnesses.

Caption: this is the very best photo of the South Carolina Lizardman, though sadly I couldn’t find the version with the top hat and cane. Taken by a mysteriously anonymous source.

Also worth noting is that the 1950s were the time when amphibious fish-monster creatures were being depicted on the big screen. The sensational and highly popular Creature from the Black Lagoon premiered in 1954 and could well have inspired people – consciously or not – to think or pretend that they might really encounter ‘frog people’ or ‘lizard people’ of this sort.

Caption: is it really coincidental that the Creature from the Black Lagoon appeared the year prior to the first appearance of the Loveland Frog? Well, probably not. Image: public domain (original here).

One more thing. If we’re going to take seriously the idea that Hunnicut and the other alleged witnesses saw real animals and misidentified them, the possibility that they saw big reptiles has to be considered quite unlikely, as noted above. Deer, standing at the roadside and seen in front view, sometimes look like humanoid bipeds since their bodies, hindlimbs and snouts merge into invisibility. I came up with this idea myself after seeing a scary roadside ‘biped’ with a round body, wide neck and slender legs morph into a deer as the car I was in approached and passed it, and I think that a few very odd sightings of similar creatures (like John Irwin’s Wharton State Forest monster of December 1993; Coleman 1995) could be explained the same way. Could confused observations of this sort explain creatures like the Loveland Frog? It’s worth considering.

Caption: how do we explain (or attempt to explain) ‘monster’ sightings like the creature reported by John Irwin in 1993? Irwin’s drawing (from Coleman 1995) is shown at left. The ‘monster’ here has several deer-like features. Could it be that Irwin saw a foreshortened deer and misinterpreted it as a biped? The deer image at right (a Wapiti female) is from Geist (1999). Images: Coleman (1995), Geist (1999).

It came from the Squamozoic. The existence of metre-tall, bipedal reptile-like creatures has to be regarded as fairly unlikely, especially when those reptile-like creature are seen carrying mechanical devices that emit sparks. The real identity behind the creature is obvious, but only if we admit the reality of parallel universes, time travel, and the ability of some creatures to somehow move between disparate points in space and time.

The Loveland Frog was no giant, humanoid frog at all, but actually a giant, vaguely humanoid lizard, and specifically one of the short-faced, big-brained iguanians from the parallel Earth of the Squamozoic.

Caption: short-faced, tailless, bipedal body forms evolved on a few occasions among the iguanians of the Squamozoic, most famously in the terrameleons (this is a Terrible terrameleon). It’s not a big step from here to time-travelling, intelligent, tool-using iguanians in 20th century Ohio. Image: electriceel.

On Squamozoic Earth, squamates (lizards, snakes and amphisbaenians) evolved to dominate the large-bodied animal fauna of the planet, and heightened intelligence evolved on several occasions. We conclude that the Loveland Frog is one of the intelligent American iguanians, presumably one that comes from a point in time somewhere in the future relative to our own position in the timeline. Whether these intelligent, parallel-universe iguanians have learnt to master time-travel and hence are deliberately travelling to 20th century Ohio as part of an exploratory or invasive mission, or whether they are merely falling inadvertently through some sort of interdimensional window, we cannot know, but perhaps we will in time.

Caption: a large, intelligent iguanian from the Squamozoic surely explains the Loveland Frog (and likely Lizardman and similar cryptids too). Here are but a few of the Squamozoic’s many denizens. Image: Darren Naish.

Our conclusion that some mystery creatures encountered on Earth are actually travellers from parallel dimensions was unashamedly inspired by promotion of the same idea, presented as a serious possibility in some of the mystery animal literature (Keel 1975, Bord & Bord 1980) and clearly not contradicted by our understanding of the way reality works.

Caption: the Bords were surely right, and John Keel was too.

On that note, The Cryptozoologicon Volume 2 will appear one day, we promise.

For previous TetZoo articles on the Cryptozoologicon project and on cryptozoology and mystery creatures in general, see…

• Tales from the Cryptozoologicon: the Yeti, August 2013

• Tales from the Cryptozoologicon: Megalodon!, August 2013

• The Cryptozoologicon (Volume I): here, at last, December 2013

• If Bigfoot Were Real, June 2016

• Bigfoot’s Genitals: What Do We Know?, August 2018

• A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, November 2019

The publication of The Cryptozoologicon Volume 2 and other various in-prep TetZoo projects is in part contingent on crowd-funded support, as is the continued appearance of new articles at this blog. Huge thanks to those who support TetZoo at patreon. If you’re interested in pledging support and seeing in-prep work, please click here.

Refs - -

Bord, J. & Bord, C. 1980. Alien Animals. Granada, London.

Bord, J. & Bord, C. 1989. Modern Mysteries of the World. Guild Publishing, London.

Coleman, L. 1995. Jersey Devil walks again. Fortean Times 83, 49.

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

Geist, V. 1999. Deer of the World. Swan Hill Press. Shrewsbury.

Haupt, R. 2015. The Loveland Frog. Skeptoid Podcast. Skeptoid Media, 30 Jun 2015. Web. 11 Jan 2020.

Keel, J. 1975. Strange Creatures from Time & Space. Nevill Spearman, London.

Naish, D. 2017. Hunting Monsters. Arcturus, London.

Newton, M. 2005. Encyclopedia of Cryptozoology. McFarland & Company, Jefferson (N. Carolina) and London.

Shuker, K. P. N. 2008. Dr Shuker’s Casebook: In Pursuit of Marvels and Mysteries. CFZ Press, Woolsery (Devon, UK).