The New World Leaf-Nosed Bat Radiation

I’ve said a few times here at TetZoo that bats have never really been given adequate coverage. This isn’t because I’m not interested in them: on the contrary, I think about bats more than I think about most other groups of mammals, and I see them and watch them more often than I do most other mammal groups. For a group that includes about 18% of extant mammalian species (using 2019 figures*), I can’t pretend to have ever given bats fair coverage. Having said all that, bats have actually been covered at TetZoo a fair bit: there was an entire 20-part series on vesper bats (properly Vespertilionidae) at ver 3, and I also published several ver 2 articles on the history and evolution of vampire bats, and on much else besides. The fact that all of these articles have been rendered worthless via the removal of their images is mightily dispiriting though, and essentially means that I need to start from scratch.

* c 6495 mammal species, c 1200 bat species.

TetZoo Towers bat library. The several boxfiles of reprints and photocopied articles are not shown. Image: Darren Naish.

TetZoo Towers bat library. The several boxfiles of reprints and photocopied articles are not shown. Image: Darren Naish.

Here, I want to talk about a group I don’t think I’ve ever covered at TetZoo before, namely the phyllostomids, or New World leaf-nosed bats, American leaf-nosed bats or spear-nosed bats. This is a large, American group that contains around 200 living species, making it the third largest bat family (vesper bats are the biggest group, followed by fruit bats). The group has sometimes been called Phyllostomatidae – the vernacular version of which is phyllostomatid – but this is less popular than Phyllostomidae. I have no idea which is really correct here and opt to merely follow majority usage on these sorts of things (insert quote from Gene Gaffney**). It’s not strictly true that I’ve never covered phyllostomids before, since vampires – once upon a time given their own eponymous family (Desmodontidae) – are now universally agreed to be nested within Phyllostomidae, and I have at least written about them.

Chrotopterus , a big spear-nosed bat. Notice how this bat has relatively broad, low-aspect wings and a large, deep uropatagium (the membrane between the legs). Contrast this with some of the images below. Image: George Henry Ford, public domain (original   here  ).

Chrotopterus, a big spear-nosed bat. Notice how this bat has relatively broad, low-aspect wings and a large, deep uropatagium (the membrane between the legs). Contrast this with some of the images below. Image: George Henry Ford, public domain (original here).

Phyllostomids occur from Argentina in the south to the southern USA (Nevada being their most northerly occurrence) in the north, and they’re highly diverse ecologically and behaviourally. They include insectivores, frugivores, nectarivores, palynivores (that’s pollen-eaters), omnivores, animalivores and (of course) obligate sanguivores. Numerous different taxonomic subdivisions have been named. We don’t need to worry about any of this in detail but, in simplified terms, Macrotinae (big-eared bats), Micronycterinae (little big-eared bats) and Desmodontinae (vampires) are outside a much larger clade that includes Vampyrinae (false vampires and kin) and Phyllostominae (spear-nosed bats and kin) as well as the nectarivorous and frugivorous Glossophaginae (long-tongued and long-nosed bats) and Stenodermatinae (American fruit bats, fig-eating bats and kin) (Baker et al. 1989, 2003, 2012; but see Wetterer et al. 2000). Vampyrinae is a clade within Phyllostominae according to some studies, in which case it gets down-graded to Vampyrini (Baker et al. 2003). All of this is depicted in a cladogram below.

Some phyllostomid portraits. At left: Big-eared woolly bat or Peter’s false vampire  Chrotopterus auritus . At right: Hairy big-eyed bat  Chiroderma villosum . Images: both Guilherme Garbino, wikipedia, CC BY-SA 4.0 (originals   here   and   here  ).

Some phyllostomid portraits. At left: Big-eared woolly bat or Peter’s false vampire Chrotopterus auritus. At right: Hairy big-eyed bat Chiroderma villosum. Images: both Guilherme Garbino, wikipedia, CC BY-SA 4.0 (originals here and here).

Phyllostomids are mostly brownish bats with simple, narrow ears. A nose-leaf – typically simple and spear-shaped – is common but not present in all species, a tragus is always present, and the species that lack nose-leaves have chin-leaves (or a series of chin ‘warts’) instead. Facial stripes are common, dark dorsal stripes are present in a few species, and such things as white patches at the wing tips and yellow rims to the ears and nose-leaves are present in some (Hill & Smith 1984). The tail is variously long, or short, and even absent altogether in some taxa, and similar variation is present in the uropatagium, or tail membrane.

The tail and uropatagia (the membranes joining the inner sides of the legs to the tail) are reduced, and sometimes highly reduced, in some phyllostomids. Here, we see this reduced condition in (at left) the Toltect fruit-eating bat  Dermanura tolteca  and (at right) in a Little yellow-shouldered bat  Sturnira lilium . Images: M.H. de Saussure, 1860, in public domain (original   here  ); Tobusaru, wikipedia CC BY 3.0 (original   here  ).

The tail and uropatagia (the membranes joining the inner sides of the legs to the tail) are reduced, and sometimes highly reduced, in some phyllostomids. Here, we see this reduced condition in (at left) the Toltect fruit-eating bat Dermanura tolteca and (at right) in a Little yellow-shouldered bat Sturnira lilium. Images: M.H. de Saussure, 1860, in public domain (original here); Tobusaru, wikipedia CC BY 3.0 (original here).

Skeletally, phyllostomids are robust, have a distinctive humerus where the distal end is angled relative to the shaft, and have a prominent secondary articulation between the large bony lump (properly termed the greater tuberosity) at the proximal end of the humerus and the scapula (Czaplewski et al. 2007). That’s right: a number of bat groups have an accessory peg-in-socket articulation involving the humerus and the body of the scapula. This means that the humerus and scapula are locked together during the upper part of the wing stroke (Hill & Smith 1984).

The most prominent exception to the ‘phyllostomids are mostly brown’ generalisation is the Honduran white bat  Ectophylla alba , sometimes likened to a fuzzy ping-pong ball and well known for its habit of constructing tents by biting through leaf ribs such that the two sides of the leaf droop on either side of the central axis. Note the yellow ears and nose leaf! The individual at left is releasing a bit of urine. Images:   Geoff Gallice  , wikipedia, CC BY 2.0 (original   here  );   Leyo  , wikipedia, CC BY-SA 2.5. (original   here  )

The most prominent exception to the ‘phyllostomids are mostly brown’ generalisation is the Honduran white bat Ectophylla alba, sometimes likened to a fuzzy ping-pong ball and well known for its habit of constructing tents by biting through leaf ribs such that the two sides of the leaf droop on either side of the central axis. Note the yellow ears and nose leaf! The individual at left is releasing a bit of urine. Images: Geoff Gallice, wikipedia, CC BY 2.0 (original here); Leyo, wikipedia, CC BY-SA 2.5. (original here)

Some phyllostomids are really exceptional as goes their anatomical and behavioural novelty. Perhaps the most remarkable are the long-tongued glossophagine flower bats, some of which have extraordinary tubular snouts, remarkably long tongues tipped with papillae, and a highly reduced dentition. The most extreme example of this sort of thing is the Banana bat, Trumpet-nosed bat or Colima long-nosed bat Musonycteris harrisoni of Mexico, an ‘extreme’ mammal as goes snout length. It’s fairly typical for people who aren’t that familiar with bat diversity to confuse glossophagines with the Old World flower-feeding megabats grouped together in Macroglossinae. There’s obviously a degree of evolutionary convergence here, though it hasn’t been that well explored in the literature, to my knowledge. Various glossophagines have symbiotic relationships with sympatric plants. Incidentally, Pallas’s long-tongued bat Glossophaga soricina is able to see UV light (Winter et al. 2003).

Some distantly related (but broadly similar) members of the phyllostomid clade Glossophaginae. At left: long-tongued champion, Pallas’s long-tongued bat  Glossophaga soricina . At right: Underwood’s long-tongued bat  Hylonycteris underwoodii . Images: Betty Wills, wikipedia CC BY-SA 4.0 (original   here  ); Karin Schneeberger/  Felineora  , wikipedia CC BY-SA 3.0 (original   here  ).

Some distantly related (but broadly similar) members of the phyllostomid clade Glossophaginae. At left: long-tongued champion, Pallas’s long-tongued bat Glossophaga soricina. At right: Underwood’s long-tongued bat Hylonycteris underwoodii. Images: Betty Wills, wikipedia CC BY-SA 4.0 (original here); Karin Schneeberger/Felineora, wikipedia CC BY-SA 3.0 (original here).

Entirely different specialisations are seen in the short-faced, frugivorous phyllostomids included within Stenodermatinae. These have flattened, broad teeth, typically have white facial stripes (an aposematic warning of their powerful bites?), and are sometimes handsome or even cute, big-eyed bats. One of the strangest of bats – the Wrinkle-faced or Lattice-winged bat Centurio senex – belongs to this group. The naked, wrinkled faces of males are mostly concealed by massive skin flaps when the bat is roosting or sleeping. There are also neck glands that seem to secrete scent, and obvious transverse bands on the wing membranes.

Resting Wrinkle-faced bats  Centurio senex  partially conceal their faces beneath thick skin folds. Translucent patches on the lower of these skin folds seem to allow these bats to detect light-level changes even when their faces are covered. Image: Jplevraud, wikipedia CC BY-SA 3.0 (original   here  ).

Resting Wrinkle-faced bats Centurio senex partially conceal their faces beneath thick skin folds. Translucent patches on the lower of these skin folds seem to allow these bats to detect light-level changes even when their faces are covered. Image: Jplevraud, wikipedia CC BY-SA 3.0 (original here).

 My favourite phyllostomids are very different from tubular-snouted flower-feeders and short-face fruit-eaters: they are the robust, more generalised species traditionally lumped together in Phyllostominae (though the name Vampyrinae has also been used for some of them). These are mostly omnivores that eat insects, fruit and small vertebrates, and some are specialised predator bats that variously catch and eat amphibians, mammals (including other bats) and birds. They include Peters’s woolly false vampire Chrotopterus auritus, the Frog-eating bat Trachops cirrhosus – famous for eating frogs and selecting them on the basis of their calls – and the spectacular Linnaeus’s false vampire Vampyrum spectrum, a predatory giant that can, in cases, have a wingspan of over 1 meter.

Vampyrum , the False vampire, has to be considered one of the most awesome of all bats. It’s convergently similar to the distantly related megadermatid bats of Africa, Asia and Australasia, also (confusingly) often called false vampires. Image: Marco Tschapka, wikipedia, CC BY-SA 3.0 (original   here  ).

Vampyrum, the False vampire, has to be considered one of the most awesome of all bats. It’s convergently similar to the distantly related megadermatid bats of Africa, Asia and Australasia, also (confusingly) often called false vampires. Image: Marco Tschapka, wikipedia, CC BY-SA 3.0 (original here).

The very impressive skull of  Vampyrum . It is robust, with big, strong teeth, especially prominent upper canines (which have an additional internal cusp) and a prominent sagittal crest. The skull can be 5.1 cm long in total (which is big for a bat). Image: Naturalis Biodiversity Center, wikipedia, public domain (original   here  ).

The very impressive skull of Vampyrum. It is robust, with big, strong teeth, especially prominent upper canines (which have an additional internal cusp) and a prominent sagittal crest. The skull can be 5.1 cm long in total (which is big for a bat). Image: Naturalis Biodiversity Center, wikipedia, public domain (original here).

When this variation in feeding ecology is mapped onto a phylogeny, it would appear that the earliest phyllostomids were insectivorous, that omnivory, nectarivory (or nectivory, take your pick) and palynivory evolved from among these insectivores, and that frugivores evolved from among nectarivores and palynivores (Baker et al. 2012). The highly specialised vampires appear – according to phylogenetic data – to have evolved directly from insectivores (which is a surprise in view of some models proposed to explain vampire evolution) and at least some members of the main frugivorous clade appear to have reverted to insectivory (Baker et al. 2012; but see Wetterer et al. 2000). Of the various evolutionary events that must have occurred here, it’s the transition to obligate frugivory that seems to have been the most successful, since the frugivorous clade is the largest (as in, most species-rich) within Phyllostomidae, containing about 70 species in 20 genera.

A few more vertebrate-eating phyllostomids. At left: California leaf-nosed bat  Macrotus californicus , the most northerly occurring phyllostomid. At right: Fringe-lipped bat  Trachops cirrhosus , a widespread species of Central and South America that eats seeds, fruits, arthropods and lizards in addition to frogs. Images: National Wildlife Service, wikipedia, public domain (original   here  ); Karin Schneeberger/  Felineora  , wikipedia CC BY 3.0 (original   here  ).

A few more vertebrate-eating phyllostomids. At left: California leaf-nosed bat Macrotus californicus, the most northerly occurring phyllostomid. At right: Fringe-lipped bat Trachops cirrhosus, a widespread species of Central and South America that eats seeds, fruits, arthropods and lizards in addition to frogs. Images: National Wildlife Service, wikipedia, public domain (original here); Karin Schneeberger/Felineora, wikipedia CC BY 3.0 (original here).

This is also the radiation that’s seemingly resulted in the greatest, most rapidly evolved amount of morphological variation, since everything here seems to have happened within the last 10 million years and has given rise to taxa that are among the most divergent and specialised of phyllostomids. Also of interest here is that the frugivorous clade appears to have evolved in the Antilles before invading the mainland (Dávalos 2007), a case of ‘upstream colonisation’ that contradicts traditional scenarios whereby continental animals give rise (via ‘downstream colonisation’) to island-dwelling forms.

Substantially simplified phyllostomid cladogram, based mostly on Baker  et al . (2003), and using their nomenclature (though they regarded false vampires - as Vampyrini - as nested within Phyllostominae). Images (top to bottom):  Macrotus  = National Wildlife Service, wikipedia, public domain (original   here  );  Desmodus  = Uwe Schmidt, wikipedia, CC BY-SA 4.0 (original   here  );  Vampyrum  = Marco Tschapka, wikipedia, CC BY-SA 3.0 (original   here  );  Phyllostomus  = Karin Schneeberger/  Felineora  , wikipedia, CC BY 3.0 (original   here  );  Platalina  = Juan A. Malo de Molina, wikipedia, CC BY-SA 3.0 (original   here  );  Sturnira  = Burtonlim, wikipedia, CC BY-SA 3.0 (original   here  ).

Substantially simplified phyllostomid cladogram, based mostly on Baker et al. (2003), and using their nomenclature (though they regarded false vampires - as Vampyrini - as nested within Phyllostominae). Images (top to bottom): Macrotus = National Wildlife Service, wikipedia, public domain (original here); Desmodus = Uwe Schmidt, wikipedia, CC BY-SA 4.0 (original here); Vampyrum = Marco Tschapka, wikipedia, CC BY-SA 3.0 (original here); Phyllostomus = Karin Schneeberger/Felineora, wikipedia, CC BY 3.0 (original here); Platalina = Juan A. Malo de Molina, wikipedia, CC BY-SA 3.0 (original here); Sturnira = Burtonlim, wikipedia, CC BY-SA 3.0 (original here).

Where in the bat tree? What sort of bats are phyllostomids, and what do we know about their evolutionary history? On the basis of anatomical characters, bat experts have generally thought that phyllostomids are close allies of naked-backed, moustached or ghost-faced bats (Mormoopidae) and bulldog bats and kin (Noctilionidae), the whole lot being grouped together in a clade termed either Phyllostomatoidea or Noctilionoidea (and it’s the last of those terms that should be preferred, so I understand). In turn, this group was thought – again, on the basis of anatomical characters – to be closely related both to vesper bats and their kin (Vespertilionoidea), and to a clade that includes both sheath-tailed bats and kin (Emballonuroidea) and horseshoe bats and kin (Rhinolophoidea) (Smith 1976).

Prior to recent (post-2000-ish) molecular studies, noctilionoids were thought to be close kin of rhinolophoids as well as emballonuroids and vespertilionoids. Rhinolophoids are now known to belong elsewhere. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project,   progress on which can be seen here.   Image: Darren Naish.

Prior to recent (post-2000-ish) molecular studies, noctilionoids were thought to be close kin of rhinolophoids as well as emballonuroids and vespertilionoids. Rhinolophoids are now known to belong elsewhere. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project, progress on which can be seen here. Image: Darren Naish.

Molecular studies, mostly published since 2000, have substantially revised our view of the bat family tree, however, and it’s now clear that rhinolophoids are not close to the other groups listed here at all (they are, instead, close relatives of megabats). Noctilionoids are still close kin of vespertilionoids, however. It also now seems that Mystacinidae and Myzopodidae are part of Noctilionoidea (Jones et al. 2002, 2005, Teeling et al. 2005, 2012). I’ll be talking more about ideas on bat phylogeny in a future article.

Simplified cladogram depicting the affinities of several of the bat groups shown - via morphological and molecular studies - to belong together within Noctilionoidea. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project,   progress on which can be seen here.   The  Vampyrum  representing Phyllostomidae, incidentally, is a placeholder which needs replacing (the existing illustration was copied directly from the work of another artist). Image: Darren Naish.

Simplified cladogram depicting the affinities of several of the bat groups shown - via morphological and molecular studies - to belong together within Noctilionoidea. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project, progress on which can be seen here. The Vampyrum representing Phyllostomidae, incidentally, is a placeholder which needs replacing (the existing illustration was copied directly from the work of another artist). Image: Darren Naish.

What does the fossil record say about phyllostomid history? The pre-Pleistocene phyllostomid record is not great but it’s still at least good enough to show that the extinct phyllostomids of the Miocene – most notably those from La Venta in Colombia – were superficially much like living ones, and that the extinct species concerned were doing the sorts of things that phyllostomids do today. The group had almost certainly, therefore, undergone its main flowering and diversification by around 20 million year ago. The Pleistocene phyllostomid record, in contrast, is good and numerous extant taxa are known from sediments of this age. 

Beautiful illustration of Salvin’s big-eyed bat  Chiroderma salvini , a stenodermatine phyllostomid that has a wide range across South and Central America. The facial stripes are not normally this pronounced in life, though it should be noted that populations are variable as goes stripe thickness. Image: Joseph Smit, in public domain (original   here  ).

Beautiful illustration of Salvin’s big-eyed bat Chiroderma salvini, a stenodermatine phyllostomid that has a wide range across South and Central America. The facial stripes are not normally this pronounced in life, though it should be noted that populations are variable as goes stripe thickness. Image: Joseph Smit, in public domain (original here).

And that’s where we’ll end things for now. I’d like to say a lot more about these bats, so we’ll be returning to them in time. And in fact I need to say a lot more about bats in general, so stay tuned for that too.

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!

For previous TetZoo articles on bats (concentrating here on articles that haven’t been stripped of images, as is the case for all ver 2 articles and the vast majority of ver 3 articles)…

Refs - -

Baker, R. J., Bininda-Emonds, O. R. P., Mantilla-Meluk, H., Porter, C. A. & Van Den Bussche, R. A. 2012. Molecular time scale of diversification of feeding strategy and morphology in New World leaf-nosed bats (Phyllostomidae): a phylogenetic perspective. In Gunnell, G. & Simmons, N. (eds). Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Cambridge, pp. 385-409.

Baker, R. J., Hoofer, S. R., Porter, C. A. & Van Den Bussche, R. A. 2003. Diversification among New World leaf-nosed bats: An evolutionary hypothesis and classification inferred from digenomic congruence of DNA sequence. Occasional Papers, Museum of Texas Tech University 230, 1-32.

Baker, R. J., Hood, C. S. & Honeycutt, R. L. 1989. Phylogenetic relationships and classification of the higher categories of the New World bat family Phyllostomidae. Systematic Zoology 38, 228-238.

Czaplewski, N. J. 1997. Chiroptera. In Kay, R. F., Madden, R. H., Cifelli, R. L. & Flynn, J. J. (eds) Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press (Washington and London), pp. 410-431.

Dávalos, L. M. 2007. Short-faced bats (Phyllostomidae: Stenodermatinae): a Caribbean radiation of strict frugivores. Journal of Biogeography 34, 364-375.

Hill, J. E. & Smith, J. D. 1984. Bats: A Natural History. British Museum (Natural History), London.

Jones, K. E., Bininda-Emonds, O. R. P. & Gittleman, J. L. 2005. Bats, clocks, and rocks: diversification patterns in Chiroptera. Evolution 59, 2243-2255.

Jones, K. E., Purvis, A., MacLarnon, A., Bininda-Emonds, O. R. P. & Simmons, N. B. 2002. A phylogenetic supertree of the bats (Mammalia: Chiroptera). Biological Reviews 77, 223-259.

Smith, J. D. 1976. Chiropteran Evolution. Texas Tech University, Lubbock.

Teeling, E. C., Dool, S. & Springer, M. S. 2012. Phylogenies, fossils and functional genes: the evolution of echolocation in bats. In Gunnell, G. & Simmons, N. (eds). Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Cambridge, pp. 1-22.

Teeling, E. C., Springer, M. S., Madsen, O., Bates, P., O’Brien, P. & Murphy, W. J. 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307, 580-584.

Wetterer, A. L., Rockman, M. V. & Simmons, N. B. 2000. Phylogeny of phyllostomid bats (Mammalia: Chiroptera) data from diverse morphological systems, sex chromosomes, and restriction sites. Bulletin of the American Museum of Natural History 248, 1-200.

Winter, Y., López, J. & von Helversen, O. 2003. Ultraviolet vision in a bat. Nature 425, 612-614.

** In a technical article on fossil side-necked turtles, Gaffney said of a very similar nomenclatural disagreement: “it’s true, I don’t give a rat’s ass which is used”.

Sleep Behaviour and Sleep Postures in Non-Human Animals

Long-time readers of TetZoo – those who’ve been visiting since the ver 2, or ScienceBlogs, years – might remember the article on sleep behaviour and sleep postures, published way back in September 2008 (that original article is here). I don’t want to get into the habit of republishing old stuff ‘from the archives’ here at ver 4, partly because I have an essentially infinite amount of new stuff I want to get online but also because the old stuff needs to be published in book form (yeah… how’s it going with that? Not well).

The sleeping behaviour of some animals is quite familiar to us. Other animals? — less so. Images: Malcolm Daniel, used with permission (left), Darren Naish (right).

The sleeping behaviour of some animals is quite familiar to us. Other animals? — less so. Images: Malcolm Daniel, used with permission (left), Darren Naish (right).

Anyway, the sleep article is sufficiently useful that it needs to be republished, especially so given that hosting issues at ScienceBlogs mean that all my old articles have been stripped of their images, sigh. Pre-amble done, let’s get to it. I’ve augmented and expanded the original text in view of new things I’ve learnt and new studies I’ve been made aware of.

Some sort of bear-like creature, asleep. Image: Darren Naish.

Some sort of bear-like creature, asleep. Image: Darren Naish.

For a long time now I have been, shall we say, gently encouraged by two of my friends to write about a subject that is both familiar, and yet also strangely alien and poorly understood. Sleep behaviour. We still know comparatively little about this subject: not only about the big stuff like its function, but even about its distribution within animals. I am not, by the way, about to tackle the big questions about sleep, nor am I going to discuss the different types of sleep (e.g., REM vs NREM sleep) and on how they differ from creature to creature. Instead I’m interested in the more superficial stuff, like how and where animals sleep, and on the postures they adopt.

John Conway’s sleeping  Tyrannosaurus , from    All Yesterdays   . We don’t know how non-bird dinosaurs slept (though several fossils give us a good idea for some groups at least). We can, nevertheless, make lots of inferences based on the behaviour of living animals. Image: John Conway/   All Yesterdays   .

John Conway’s sleeping Tyrannosaurus, from All Yesterdays. We don’t know how non-bird dinosaurs slept (though several fossils give us a good idea for some groups at least). We can, nevertheless, make lots of inferences based on the behaviour of living animals. Image: John Conway/All Yesterdays.

What we do know about sleep is scattered widely in the literature and it isn’t easy to obtain any sort of review of the subject. A chapter in an academic book – Lee-Chiong’s 2006. Sleep: A Comprehensive Handbook – does include a good overview of sleep behaviour in non-human animals (Lee-Chiong 2006, Lesku et al. 2006), but the google books version (the only one I’ve seen) doesn’t include the chapter in entirety, as per usual. There is at least one book devoted to sleep in non-human aniamls: Maurice Burton’s Sleep and Hibernation in the Animal World (Odhams Books, 1969), one of twelve books written for the Animal World in Colour series. Burton is well known for amassing large amounts of anecdotal data on the natural world (he also wrote books covering cryptozoology and the emotional lives of animals), and while the books in the Animal World in Colour series include a lot of often bizarre and fascinating information, they’re completely unreferenced and at least some of the content looks apocryphal and difficult to be confident about. Despite these problems I’ve borrowed heavily from his book on sleep here.

Maurice Burton’s  Sleep and Hibernation in the Animal World , part of the Animal World in Colour series, shown at left at their home in the TetZoo library (there are 12 titles in total, so I’m still missing three). Images: Darren Naish.

Maurice Burton’s Sleep and Hibernation in the Animal World, part of the Animal World in Colour series, shown at left at their home in the TetZoo library (there are 12 titles in total, so I’m still missing three). Images: Darren Naish.

Needless to say, I’m only going to be looking here at sleep in tetrapods, though where relevant I will mention other animals. The first thing to say is that some animals get by with very little sleep, or in fact may even go without sleep entirely. Among vertebrates, all identified non-sleeping taxa are fish (Kavanau 1998). The species concerned have little in common (they include blind cave-dwellers, various sharks including some triakids, lamnids, hammerheads and sevengills, and mackerels and scombrids), but all have lifestyles where “visual input is greatly reduced or absent during lengthy periods”, where “schooling greatly reduces needs for sensory information, particularly visual'” or where the animals lead a “comparatively routine existence in essentially featureless, open waters” (Kavanau 1998, p. 269).

Fishes. I ain’t sayin’ nothing, apart from what I just said. Image: Darren Naish.

Fishes. I ain’t sayin’ nothing, apart from what I just said. Image: Darren Naish.

Reduced or delayed sleep. Perhaps because no tetrapod lives in this way, it seems that all indulge in at least some sleep activity, and indeed will die if deprived of sleep for prolonged periods (Rechtschaffen et al. 1983). Having said that, some tetrapods sleep very little, or go for comparatively long periods with little to no sleep. One proposal for the function of sleep is that it gives animals the chance to save energy when remaining active is unproductive. Within this model, animals might be able to forego sleep if remaining active is productive, as would be the case during periods when an otherwise rare food source is suddenly in plentiful supply, during migrations, or during a brief breeding season when wakefulness guarantees successful mating.

On that last point, male Pectoral sandpipers Calidris melanotos are known to reduce their time spent sleeping during their annual lekking period, those males that sleep the least being the ones that succeed in having the highest number of copulations (Lesku et al. 2012). In other words, sexual selection has led to a reduction in sleep. We might honestly consider whether this is applicable to other animals that don’t sleep as much or as often as expected, our own species included (read on).

Pectoral sandpiper, the most wideawake of the waders… pending future discoveries. Image:  Andy Reago & Chrissy McClarren , wikipedia, CC BY 2.0 (original  here ).

Pectoral sandpiper, the most wideawake of the waders… pending future discoveries. Image: Andy Reago & Chrissy McClarren, wikipedia, CC BY 2.0 (original here).

The animals best known for reducing or delaying their sleep are seabirds. Some species are on the wing for weeks, months and supposedly even years at a time and appear to be constantly active. Yes, years at a time. Couzens (2008) writes of Sooty terns Onychoprion fuscata staying aloft for as much as four or five years as juveniles and that “Nobody has ever seen a healthy juvenile bird at rest, whether on a beach, rock, buoy or on any floating object” and “The thought of keeping aloft for four or more years is mind-boggling to us. How do the birds sleep? Why don’t they tire? What about storms?” (p. 83).

It has been suggested that such birds might sleep on the wing – this has so far been confirmed for frigatebirds using electroencephalograms and GPS loggers (Rattenborg et al. 2016) – but it’s considered equally plausible that they simply delay their sleep to some point later in life. And having mentioned ‘sleep on the wing’, while it’s well known that swifts sleep in flight, the evidence for this (which essentially consists of observations of swifts seen decreasing in altitude during the early hours of the morning) is poor and we still require detailed studies (Rattenborg 2006).

I’ve seen hundreds of frigatebirds while in South America - this is my best photo of one - but I don’t remember noticing that any were asleep. Image: Darren Naish.

I’ve seen hundreds of frigatebirds while in South America - this is my best photo of one - but I don’t remember noticing that any were asleep. Image: Darren Naish.

Cetaceans – or, killer whales and bottlenose dolphins at least – go for the first month or two of life without sleeping, and their accompanying mothers also go without sleep during this period (Lyamin et al. 2005). The possibility exists that very short bursts of shallow sleep might have occurred in the animals under study, but any such bursts (which weren’t detected) could not have exceeded 30 seconds in duration (Lyamin et al. 2005). Quite how these animals can go without proper sleep during a period coinciding with important events in growth and development is unknown but it’s consistent with the idea that sleep can be dispensed with “when ecological demands favour wakefulness” (Lesku et al. 2012, p. 1654).

Sleeping cow, photographed in India, in a pose not considered that typical for bovids or artiodactyls in general. It’s not  that  uncommon if you look at sufficient numbers of sleeping cows, however. Image: the original source has disappeared from the internet!

Sleeping cow, photographed in India, in a pose not considered that typical for bovids or artiodactyls in general. It’s not that uncommon if you look at sufficient numbers of sleeping cows, however. Image: the original source has disappeared from the internet!

Some ruminants seem to engage in very little sleep. This might be because they need to remain constantly alert for predators, but it might also be because their biology requires a virtually continuous cycle of eating and ruminating. Burton (1969) wrote of observations made on goats which suggest that, during a rest period that lasts about 8 hours, they never really relax fully, and it’s even stated here and there that some ruminants (like cattle) sleep with their eyes open. Ruminants do sleep with their eyes closed at least sometimes however, as verified by the sleeping cow shown above. Passive stay apparatus (special locking mechanisms in the limbs) mean that some hoofed mammals can remain standing virtually indefinitely, and such mammals (which include deer, rhinos and horses) are therefore capable of sleeping while standing. Having said that, they will still lie down to rest when they want to. I photographed the donkey shown here as I initially thought it was dead.

Sleeping domestic donkey, reclining on its side. The animal stayed like this for some time (an hour at least). Image: Darren Naish.

Sleeping domestic donkey, reclining on its side. The animal stayed like this for some time (an hour at least). Image: Darren Naish.

Finally, also worth noting here is that humans actually sleep way less than we ‘should’, relative to other primates. For whatever reason, we’ve managed to reduce the amount of sleep we should have according to our body size, perhaps by as much as 50%. I take seriously the possibility that we’re similar to Pectoral sandpipers, and that sleep reduction is driven by sexual selection (this is the get lucky hypothesis). Other possibilities have equally good explanatory power though. A whole article could be written on human sleep behaviour…

Human sleep behaviour is interesting - we’re unusual, relative to other primates, in several respects. Sleeping in the snow is not usually advised though. Image: Darren Naish.

Human sleep behaviour is interesting - we’re unusual, relative to other primates, in several respects. Sleeping in the snow is not usually advised though. Image: Darren Naish.

Giraffes and elephants. It used to be thought that the biggest extant hoofed mammals – giraffes – didn’t sleep at all, but this was shown to be incorrect by Grzimek (1956) who demonstrated that adult giraffes slept recumbently on the ground for short periods (2.5-6 minutes), resting the head on the hindquarters or ground and holding the neck in an arced posture. This posture is not unique to giraffes but is also practised by okapis and some bovids. The sleep behaviour of giraffes was looked at in detail by Tobler & Schwierin (1996) who showed that giraffes slept both while standing and while recumbent for a total of 4.6 hours per 24 hours.

Here’s a baby giraffe in the so-called ‘kettle posture’ typical for sleeping giraffes (though they can sleep in other poses too). This image is all over the internet and I haven’t succeeded in finding who should be credited for the original.

Here’s a baby giraffe in the so-called ‘kettle posture’ typical for sleeping giraffes (though they can sleep in other poses too). This image is all over the internet and I haven’t succeeded in finding who should be credited for the original.

Evidently little known is that elephants lie down on the ground to sleep properly: even today it seems widely thought that elephants ‘crush themselves under their own weight’ if they lie down, but this just isn’t true. In the wild, elephants sleep for 1-4.5 hours (per 24) in a recumbent position, with individuals showing preferences for sleeping on their left or right sides. Tobler (1992) reported that captive Asian elephants slept in a recumbent position for an average of 72 minutes a night. Elephants have been photographed propping themselves up against termite mounds and other structures when lying down to sleep, and in captivity they have been reported to construct pillows from straw. The photo below shows a sleeping captive Asian elephant Elephus maximus, and for images of African elephants sleeping recumbently in the wild go here.

Sleeping Asian elephant in captivity. Image: Fruggo, CC BY 1.0, wikipedia (original  here ).

Sleeping Asian elephant in captivity. Image: Fruggo, CC BY 1.0, wikipedia (original here).

Birds large and small, on the ground. Large birds are also known to recline on the ground for at least short periods. Burton (1969) reported how a scientist at Frankfurt Zoo monitored the sleep behaviour of ostriches: for 7-8 hours per night, they slept lightly with the neck erect. But for an average of 9 minutes each night they would lie on their sides, stretch their necks and legs out on the ground, and sleep deeply, during which time they couldn’t be roused by lights or noises (except the very loudest noises). Emus, incidentally, are said to sleep deeply with the head and neck resting on the back, rather than lying on the ground.

A Ptarmigan  Lagopus mutus  in its snow burrow. This illustration - by Ad Cameron - is meant to show the bird sleeping in its burrow, but a bird properly using such a refuge would be concealed from outside view. Image: Ad Cameron, in Perrins (1992).

A Ptarmigan Lagopus mutus in its snow burrow. This illustration - by Ad Cameron - is meant to show the bird sleeping in its burrow, but a bird properly using such a refuge would be concealed from outside view. Image: Ad Cameron, in Perrins (1992).

Moving to other birds, terrestrial, ground-foraging birds like pheasants roost on the ground or in the branches of trees, and simply huddle up and tuck in their appendages. Grouse living in snowy places construct snow burrows and have a set of adaptations that make them good at constructing these shelters (a topic I aim to cover in the near future), and various passerines of snowy environments – including kinglets, finches, buntings and tits – also build snow shelters and sleep in them (e.g., Sulkava 1969, Novikov 1972).

Left: a tree-creeper ( Certhia ), sleeping while partially tucked up inside a cavity in a tree. Right: a rendition of one of those famous cases in which large numbers of wrens ( Troglodytes troglodytes ) - more than 40 or 50 - have been discovered packed inside the same nest, huddled together for warmth. In this case, the wrens are using an empty House martin  Delichon urbicum  nest. Images: Burton (1969), Ad Cameron, in Perrins (1992).

Left: a tree-creeper (Certhia), sleeping while partially tucked up inside a cavity in a tree. Right: a rendition of one of those famous cases in which large numbers of wrens (Troglodytes troglodytes) - more than 40 or 50 - have been discovered packed inside the same nest, huddled together for warmth. In this case, the wrens are using an empty House martin Delichon urbicum nest. Images: Burton (1969), Ad Cameron, in Perrins (1992).

Small birds in cavities and thick foliage. Many small, tree-dwelling birds don’t sit out on branches during the night (though some do, see below), but secrete themselves into small crevices, cavities or thick vegetation. I once discovered a Blue tit Cyanistes caeruleus tucked deep within a crevice in the bark of a tree, only its tail tip sticking out. Small birds of many sorts do seem to rely on dense foliage as a night-time refuge. Remember this next time you unnecessarily tear down the ivy from a tree or fence.

This photo is pretty terrible but it’s the only one I have of small birds (in this case: House sparrows  Passer domesticus ) sleeping in a concealed spot, tucked up beneath a roof and perched on tangled vegetation. It was taken in May 2011 and is partly so bad because I didn’t want to disturb the birds. Image: Darren Naish.

This photo is pretty terrible but it’s the only one I have of small birds (in this case: House sparrows Passer domesticus) sleeping in a concealed spot, tucked up beneath a roof and perched on tangled vegetation. It was taken in May 2011 and is partly so bad because I didn’t want to disturb the birds. Image: Darren Naish.

Some passerines, including wrens, weavers and bush-tits, form large huddles within cavities (Perrins 1992). Burton (1969) wrote that birds which climb vertically on substrates, like nuthatches and treecreepers, also sleep in vertical postures, with nuthatches resting head-down and treecreepers resting head-up. Woodpeckers are also reported to sleep clinging vertically to vertical trunks (Perrins 1992). A few passerine species have been reported sleeping on their nests even when the nests were empty of eggs.

Trogons (a  Harpactes  species, perhaps Whitehead’s trogon  H. whiteheadi ) photographed asleep at night, on Borneo. Image: Matthew Connors, used with permission.

Trogons (a Harpactes species, perhaps Whitehead’s trogon H. whiteheadi) photographed asleep at night, on Borneo. Image: Matthew Connors, used with permission.

According to Burton (1969), birds don’t put their heads under their wings when they sleep, but instead bury the bill into the scapular feathers. He also stated that penguins are the only birds that properly hide the bill under the wing: they can’t submerge any part of their head within their scapular feathers, because their feathers are so short. I wonder what the hesperornithines did (they lacked long feathers and possessed strongly reduced wings).

Left: King penguin  Aptenodytes patagonicus  sleeping, bill tucked under wing. I can’t confirm that this is what it does show. However… Right: illustration of exactly the same thing, from Burton (1969). Bonus mystery penguin in the background. Image:  jpmatth , CC-BY-NC-ND 2.0 (original  here ), Burton (1969).

Left: King penguin Aptenodytes patagonicus sleeping, bill tucked under wing. I can’t confirm that this is what it does show. However… Right: illustration of exactly the same thing, from Burton (1969). Bonus mystery penguin in the background. Image: jpmatth, CC-BY-NC-ND 2.0 (original here), Burton (1969).

Unihemispheric sleep. So far I haven’t mentioned the fact that some animals sleep with only ‘half’ of the brain. This is called unihemispheric sleep (less frequently: unihemispherical sleep, properly unihemispheric slow-wave sleep or USWS), and it allows the animal to stay partly alert while in a restful state. Animals capable of this may be able to keep one eye open and alert for danger – this is termed unilateral eye closure or UEC – and may be able to regulate movement while resting. It’s well known for birds, including those that sleep on the wing, those that sleep on the water surface, and those that sleep in exposed terrestrial places like beaches and mudflats.

Sleeping Mute swan  Cygnus olor , not obviously employing unihemispheric sleep. Image: Neil Phillips, used with permission.

Sleeping Mute swan Cygnus olor, not obviously employing unihemispheric sleep. Image: Neil Phillips, used with permission.

Unihemispheric sleep is important to marine mammals, since they doze at or near the water surface and still need to regulate their breathing (Lyamin et al. 2008). While cetaceans typically sleep unihemispherically in a horizontal pose (this is sometimes termed ‘logging’, since a sleeping whale can look superficially like a log), some pinnipeds – including Grey seal Halichoerus grypus, Northern fur seal Callorhinus ursinus and Walrus Odobenus rosmarus – hang in a vertical posture, as do some whales, sometimes. Weddell seals Leptonychotes weddelli are also reported to do this while submerged close to breathing holes in the ice. It has also been noted that some pinnipeds sleep underwater in cetacean fashion, “rising periodically, while still asleep apparently, to take a breath and then sinking again to continue their slumber” (Burton 1969, p. 27).

A captive Harbour seal  Phoca vitulina  sleeping underwater. Image:  Rock Paper Lizard  (thanks to Vasha for the heads-up).

A captive Harbour seal Phoca vitulina sleeping underwater. Image: Rock Paper Lizard (thanks to Vasha for the heads-up).

Sleep behaviour in non-bird reptiles is not tremendously well studied and there aren’t many relevant studies out there. However, sea turtles have long been suspected to be unihemispheric sleepers. After all, they frequently sleep on the sea floor or even while resting on coral or submerged rocks. Crocodylians – caimans and crocodiles, at least – practise UEC and their behaviour during the phases when UEC is at play is similar (as goes how they respond to potential threats and so on) to that of birds and mammals that sleep unihemispherically (Kelly et al. 2015). They might, therefore, be engaging in unihemispheric sleep, though this has yet to be confirmed so far as I know.

A sleeping iguanian - an Ornate earless agama  Aphaniotis ornata , a draconine agamid endemic to Borneo - photographed at night in a forest. Image: Matthew Connors, used with permission.

A sleeping iguanian - an Ornate earless agama Aphaniotis ornata, a draconine agamid endemic to Borneo - photographed at night in a forest. Image: Matthew Connors, used with permission.

UEC is fairly well known in lizards, yet little studied. A study of Western fence lizard Sceloporus occidentalis again found that sleep behaviour during UEC – how the lizards kept open whichever eye was closest to the last observed position of a potential threat – was similar to that of birds (Mathews et al. 2006), suggesting a similar evolutionary function and that unihemispheric sleep might be at play. If crocodylians and lizards do exhibit both UEC and unihemispheric sleep, these might be ancestral for diapsids, or certainly archosaurs at least. Ergo, I’m going to promote the idea of unihemispheric sleep in non-bird dinosaurs.

Not sure if this lizard - this is a  Lacerta  species, most likely a Western green lizard  L. bilineata  - is fully asleep, but at least it has one of its eyes closed. Image: Neil Phillips, used with permission.

Not sure if this lizard - this is a Lacerta species, most likely a Western green lizard L. bilineata - is fully asleep, but at least it has one of its eyes closed. Image: Neil Phillips, used with permission.

Incidentally, REM sleep has also been reported for chameleons and in both desert iguanas (Dipsosaurus) and spiny-tailed iguanas (Ctenosaura). However, the reports are somewhat anecdotal and what’s been reported as REM sleep might instead describe observations where the animals were moving their eyes due to disturbance (Lesku et al. 2006).

That’ll do for now. This has been nothing like a thorough review of the subject and, as I said at the start, it’s been very superficial and I haven’t looked at the big questions about sleep behaviour.

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 - -

Burton, M. 1969. Sleep and Hibernation in the Animal World. Odhams Books, London.

Couzens, D. 2008. Extreme Birds. HarperCollins, London.

Grzimek, B. 1956. Schlaf von Giraffen und Okapi. Naturwissenschaften 17, 406.

Kavanau, J. L. 1998. Vertebrates that never sleep: implications for sleep's basic function. Brain Research Bulletin 46, 269-279.

Kelly, M. L., Peters, R. A., Tisdale, R. K. & Lesku, J. A. 2015. Unihemispheric sleep in crocodilians? Journal of Experimental Biology 218, 3175-3178.

Lee-Chiong, T. L. 2006. Sleep: A Comprehensive Handbook. John Wiley & Sons, Hoboken, NJ.

Lesku, J. A., Rattenborg, N. C. & Amlaner, C. J. 2006. The evolution of sleep: a phylogenetic approach. In Lee-Chiong, T. L. (ed) Sleep: A Comprehensive Handbook. John Wiley & Sons, Hoboken, NJ, pp. 49-61.

Lesku, J. A., Rattenborg, N., Valcu, M., Vyssotski, A. L., Kuhn, S., Kuemmeth, F., Heirich, W. & Kempanaers, B. 2012. Adaptive sleep loss in polygynous pectoral sandpipers. Science 337, 1654-1658.

Lyamin, O. I., Manger, P. R., Ridgway, S. H., Mukhametov, L. M. & Siegel, J. M. 2008. Cetacean sleep: an unusual form of mammalian sleep. Neuroscience and Biobehavioral Reviews 32, 1451-1484.

Lyamin, O., Pryaslova, J., Lance, V. & Siegel, J.  2005. Animal behaviour: continuous activity in cetaceans after birth. Nature 435, 1177. 

Mathews, C. G., Lesku, J. A., Lima, S. L. & Amlaner, C. J. 2006. Asynchronous eye closure as an anti-predator behavior in the western fence lizard (Sceloporus occidentalis). Ethology 112, 286-292.

Novikov, G. A. 1972. The use of under-snow refuges among small birds of the sparrow family. Aquilo, Series Zoologica 13, 95-97.

Sulkava, S. 1969. On small birds spending the night in the snow. Aquilo, Series Zoologica 7, 33-37.

Perrins, C. 1992. Bird Life: An Introduction to the World of Birds. Magna Books, Liecester [sic].

Rattenborg, N. C. 2006. Do birds sleep in flight? Naturwissenschaften 93, 413-425.

Rattenborg, N. C., Voirin, B., Cruz, S. M., Tisdale, R., Dell’Omo, D., Lipp, H.-P. Wikelski, M. & Vyssotski, A. L. 2016. Evidence that birds sleep in mid-flight. Nature Communications 7: 12468.

Rechtschaffen, A., Gilliland, M. A., Bergmann, B. M. & Winter, J. B. 1983. Physiological correlates of prolonged sleep deprivation in rats. Science 221, 182-184.

Tobler, I. 1992. Behavioral sleep in the Asian elephant in captivity. Sleep 15, 1-12.

Tobler, I. & Schwierin, B. 1996. Behavioural sleep in the giraffe (Giraffa camelopardalis) in a zoological garden. Journal of Sleep Research 5, 21-32.

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

Welcome to the second article in this short series on recently(ish) published books on the Loch Ness Monster (or LNM) (the first article is here).

The most impressive and interesting of the several Nessie paintings produced by Peter Scott - here, depicted on the back of my battered copy of the PG Tips 1987  Unexplained Mysteries of the World , written by Robert J. M. Rickard. Image: Darren Naish.

The most impressive and interesting of the several Nessie paintings produced by Peter Scott - here, depicted on the back of my battered copy of the PG Tips 1987 Unexplained Mysteries of the World, written by Robert J. M. Rickard. Image: Darren Naish.

This time round, we look at the 2015 volume A Monstrous Commotion: the Mysteries of Loch Ness, a dense, thick, attractively designed volume of 365 pages that might be the only LNM-themed book that could be classed as an airport novel (Williams 2015). I confess to being unaware of Gareth Williams prior to hearing about the publication of this book. But maybe that’s understandable, since a brief biography tells us that he’s an internationally recognised expert on diabetes and obesity affiliated with the University of Bristol, has penned over 200 papers on medical topics, and has previously published books on smallpox and polio.

Front cover of   Williams (2015)  .

Front cover of Williams (2015).

The volume begins with a timeline, a few pages providing potted biographies of the many human characters, a list of illustrations and some maps. The book also includes two plate sections and a smattering of black and white drawings.

The primary value of this book – its main selling point to an audience familiar with the LNM – is that it tells the backstory to the 1975 Nature paper by Sir Peter Scott and Robert Rines, a promise made in both the preface and the blurb on the back. This is the infamous paper – I’ll make the point again: published in the world’s most prestigious scientific journal – in which Scott and Rines argued not only that Nessie is real and that they had evidence proving it, but that it needed a scientific name. And thus we have Nessiteras rhombopteryx Scott and Rines, 1975 (and: no, it wasn’t a deliberately constructed anagram of ‘Monster Hoax by Sir Peter S’; to state such ignores Scott’s long-running, highly active investment and commitment to belief in the monster and his many published statements on it). The run-up to the publication of this paper, the fallout, and the alliances that were attempted, formed and broken is a fascinating story never told before in such depth, and it’s Williams’s use of Peter Scott’s correspondence that has allowed him to tell the tale. Excellent. This should be good.

Loch Ness is a beautiful and sublime place, monster or not. Image: Darren Naish.

Loch Ness is a beautiful and sublime place, monster or not. Image: Darren Naish.

Alas, I was immediately disappointed on finding that the book starts – as do so many books on the Loch Ness Monster – with that oh so familiar stuff about the Great Glen Fault, St Columba, kelpie legends and the stories and events of the 1930s. Clearly, we aren’t getting the Scott story alone, but the whole shebang, and while Williams writes well, I’m not sure that there’s anything in the early chapters that hasn’t been covered before.

I’ve said before that there are awful lot of books on the Loch Ness Monster… even this is far from a complete selection of what’s out there (I’m still collecting). Image: Darren Naish.

I’ve said before that there are awful lot of books on the Loch Ness Monster… even this is far from a complete selection of what’s out there (I’m still collecting). Image: Darren Naish.

His take on Rupert Gould is admittedly interesting though. Gould – typically portrayed by authors of LNM-themed books as a bold and daring adventurer, a physical and metaphorical ex-military giant of a man who covered great distances on his motorbike and was a wise and indefatigable collector of interviews and facts, and a pioneering investigator of the unknown – is portrayed as a troubled oddball deeply affected by the frightening events of warfare. And yes, Williams does cover Gould’s eventual conclusion (about-turn, if you like) that the Spicers didn’t see a giant scientifically unrecognised (semi)aquatic vertebrate species, but “a huddle of deer crossing the road” (p. 227). Incidentally, Williams states that Gould made this private admonition in the marked proofs of his book The Loch Ness Monster and Others, but that’s not right. The annotation concerned was hand-written in a published copy and not connected at all to the manuscript during its proof stage (Binns 2017, p. 150). This is one of many minor but arguably important errors made throughout the book.

Peter Macnab’s photo of 1955. This is the version lacking the vegetation in the foreground. Regarded by some Nessie proponents as depicting two monsters swimming in close proximity, it is most likely part of a boat wake, as suggested by the lines in the water about parallel to the ‘monster(s)’. This is a scan of the original photo, provided by Dick Raynor (and available  here ). Image: (c) P. A. Macnab.

Peter Macnab’s photo of 1955. This is the version lacking the vegetation in the foreground. Regarded by some Nessie proponents as depicting two monsters swimming in close proximity, it is most likely part of a boat wake, as suggested by the lines in the water about parallel to the ‘monster(s)’. This is a scan of the original photo, provided by Dick Raynor (and available here). Image: (c) P. A. Macnab.

Post-Gould, Williams discusses Constance Whyte’s More Than a Legend and the way LNM news was received at the time before going on to discuss the pop-culture backdrop to the events of the 1960s and 70s, somehow weaving in David Attenborough and Zoo Quest for a Dragon, Jacques Cousteau and Hans Hass. After a brief skip through the adventures of Torquil MacLeod and Peter Macnab (both are among those ‘classic’ eyewitnesses who claimed, respectively, a remarkable sighting and a remarkable photo… of a boat wake), we’re introduced to one of the pivotal and most influential characters in LNMology: Tim Dinsdale, aeronautical engineer, charismatic good-guy and near-professional monster believer. Dinsdale is especially relevant to the story Williams tells because it was he – not Constance Whyte, not the preponderance of evidence, not the rash of 1930s sightings – who roped Peter Scott into the saga. I don’t want to say too much about Dinsdale here since he’ll form the focus of my third LNM-themed book review.

Torquil MacLeod’s Nessie of February 1960, observed through binoculars and estimated to be 13-15 m long, as drawn by Alan Jones for   Witchell (1975)  . Image: Alan Jones/  Witchell (1975)  .

Torquil MacLeod’s Nessie of February 1960, observed through binoculars and estimated to be 13-15 m long, as drawn by Alan Jones for Witchell (1975). Image: Alan Jones/Witchell (1975).

What I will say for now is that Williams is by far too sympathetic to Dinsdale, failing to remark on Dinsdale’s sudden and, frankly, remarkable commitment to belief in the monster, nor is he appropriately critical of Dinsdale’s sightings or claims. Take Williams’s statement (p. 70) that Dinsdale was “catapulted into the limelight and would never escape from it”, or that “he found himself on Panorama, the BBC’s flagship news and current affairs programme” (p. 70). More accurate segments of text might read that Dinsdale “clamoured for and gained the limelight, and successfully managed to hold it upon himself for years to come”, and that “finally, he received the attention he had sought via a campaign of sending letters and telegrams, and succeeded in winning a spot on Panorama”.

Dinsdale (standing, at right) with Robert Rines (l) and Martin Klein in 1970. Image:   Dinsdale (1976)  .

Dinsdale (standing, at right) with Robert Rines (l) and Martin Klein in 1970. Image: Dinsdale (1976).

Dinsdale arrives. On reaching the part of the book that discusses Dinsdale and his Loch Ness adventures, I was finally feeling that I’d gotten through the preamble and reached the good stuff. To be clear, Williams does deliver the goods, providing a discussion and review novel enough and interesting enough to make the book a worthy addition to the LNM literature.

As noted above, it was Dinsdale who – through writing to Scott in a possibly desperate yet optimistic and bold gamble – brought Scott into the fold, his initial letter (addressed to ‘Mr Peter Scott, Naturalist, c/o BBC Television, London W1’) promising the support of a noted and reputable scientist (almost certainly Maurice Burton, then of the British Museum (Natural History)). Dinsdale’s several later letters discussed his mathematical analysis which basically consisted of calculating averages from the various eyewitness accounts that contained measurements.

Tim Dinsdale and his identikit-style view of what the Loch Ness Monster looked like. He reached this view by bundling all eyewitness accounts together and taking averages. Image: (c) Tim Dinsdale.

Tim Dinsdale and his identikit-style view of what the Loch Ness Monster looked like. He reached this view by bundling all eyewitness accounts together and taking averages. Image: (c) Tim Dinsdale.

Why was Scott prepared to let himself be involved in the Loch Ness story? While Scott certainly stated that his belief in the monster was based on sheer preponderance of evidence (Scott 1976), Williams notes (p. xxxiii) that Scott’s interest in Nessie was quite plausibly motivated by his feeling that it could serve as a flagship species, in the same ballpark as the tiger and giant panda, for the fledgling World Willdlife Fund. Scott’s own drawings support this idea.

Here’s another of the big, spectacular, Nessie-themed works of art produced by Peter Scott (this is only a section of the whole thing). Image: (c) Peter Scott.

Here’s another of the big, spectacular, Nessie-themed works of art produced by Peter Scott (this is only a section of the whole thing). Image: (c) Peter Scott.

Scott and Dinsdale met in person in 1960, but not before Dinsdale explained his plan. He would need Scott as an ally in convincing her majesty Queen Elizabeth II that Nessie was real and in need of protection. Scott knew the Royal Family, moved in the right circles, and was sufficiently impressed by Dinsdale’s argumentation to consider this an appropriate course of action, even making the suggestion that Nessie might be given the scientific name Elizabethia nessiae* (Williams 2015). Alas, Dinsdale had already written to the Royal Family by this time and his impetuousness on this front – he was to write to them several more times – partly derailed efforts to carefully, thoughtfully build a case for the monster’s existence, one that might be sufficiently interesting and carefully stated to keep sceptics, the scientific community, the media and people like the Royal Family on board.

* Incidentally, another proposed binomial – Nessiesaurus o’connori (sic: the specific name should have been written ‘oconnori’) – is also outed in this book. It was proposed by Peter O’Connor, author of the almost certainly hoaxed ‘inverted kayak’ photo of 1960, in his correspondence to Scott (Williams 2015).

By the mid 1970s, Peter Scott was happy to publicly state a belief in the Loch Ness Monster, and there are even photographs of him wearing an ‘I Believe in Nessie’ t-shirt. Here’s the cover of a magazine issue that features a key Scott article on the subject. Image: Darren Naish.

By the mid 1970s, Peter Scott was happy to publicly state a belief in the Loch Ness Monster, and there are even photographs of him wearing an ‘I Believe in Nessie’ t-shirt. Here’s the cover of a magazine issue that features a key Scott article on the subject. Image: Darren Naish.

Over the months and years that followed, Scott worked to build a case, Tim Dinsdale’s film of 1960 being one of several pieces of evidence deemed crucial. The many ups and downs, false-starts, setbacks, and input and involvement of others make for a complex story that I’m not about to summarise. The eventual outcome, which had emerged by 1970, was the involvement of Americans including Chicago’s Roy Mackal and patent lawyer Robert H. Rines, the rise and fall of the Loch Ness Phenomena Investigation Bureau, and a gradual parting of the ways between Scott and Dinsdale.

There’s a definite undercurrent in the book of Scott and Dinsdale working to somewhat different ends. The Dinsdale plan was to announce and promote the monster’s existence and reality as loudly and frequently as possible. The Scott plan was to form a solid portfolio of good evidence, hold formal meetings where this evidence could be presented to and digested by the right parties, and to ultimately gain legal protection for a neglected and remarkable new species honestly thought by Scott to be, most probably, a living plesiosaur (Scott 1976).

Palaeontologists specialising on plesiosaurs have near universally been very hostile to the idea that the Loch Ness Monster might be a living plesiosaur. But it’s also a very familiar idea that plesiosaur experts have sometimes toyed with. This diagram is from  Big Mouths and Long Necks , a short book devoted to plesiosaurs. Image: Taylor & Martin (1990).

Palaeontologists specialising on plesiosaurs have near universally been very hostile to the idea that the Loch Ness Monster might be a living plesiosaur. But it’s also a very familiar idea that plesiosaur experts have sometimes toyed with. This diagram is from Big Mouths and Long Necks, a short book devoted to plesiosaurs. Image: Taylor & Martin (1990).

Scott and Rines 1975, and the ‘flipper’ photos. As anyone familiar with the Loch Ness story knows, the turning point was the use of sonar detection and strobe photography in the loch, the eventual result being the presentation of photos said by Rines and his colleagues to be proof of the monster’s existence and to give insight on its form (Scott & Rines 1975, Rines et al. 1976, Scott 1976, Sitwell 1976, Rines 1982).

My own take on one of the Rines/Egerton ‘flipper photos’, drawn when I was about 14. Like many people in those years (this would’ve been drawn in the late 1980s), I had been led to believe that the photos really show the giant, diamond-shaped flippers of a very big animal. They don’t. Image: Darren Naish.

My own take on one of the Rines/Egerton ‘flipper photos’, drawn when I was about 14. Like many people in those years (this would’ve been drawn in the late 1980s), I had been led to believe that the photos really show the giant, diamond-shaped flippers of a very big animal. They don’t. Image: Darren Naish.

Initial claims that the flipper photos showed a pentadactyl anatomy (thereby confirming a tetrapod identity for the creature), that two diving animals, moving synchronously and close together, had been captured in a single frame and that a close-up view of the animal’s external surface revealed details of skin texture and even its parasites (Witchell 1975, p. 150), all proved embellished or inaccurate, to use the kindest words possible. We think today that the flipper photos were physically modified, that the ‘gargoyle head’ photo (which had been rotated by 90° relative to its original orientation) doesn’t depict an animal’s head but a tree stump on the floor of the loch, and that an alleged shot of the body and neck cannot be of a large animal but a small object close to the camera, most likely a submerged branch (Naish 2017).

The weird and ugly ‘gargoyle head’, interpreted as the snorkelled, horned, short-faced creature depicted at right in this painting by Peter Scott. Read on for another version of that Scott painting. Images: Rines  et al . (1976), Peter Scott.

The weird and ugly ‘gargoyle head’, interpreted as the snorkelled, horned, short-faced creature depicted at right in this painting by Peter Scott. Read on for another version of that Scott painting. Images: Rines et al. (1976), Peter Scott.

Nevertheless, it’s obvious from some of the things written at the time that these developments must have been extraordinarily exciting. I’m always struck by the following breathless words from Nicholas Witchell…

“This paper edition of The Loch Ness Story is being rushed out in the autumn of 1975 at a time when the world is about to witness one of the greatest and most dramatic discoveries of the twentieth century: the discovery and probable identification of a semi-mythical creature known throughout the world as the ‘Monster’ of Scotland’s Loch Ness.

“As the final chapter describes, a set of detailed colour photographs of the head and body of the ‘Monster’ have been taken by a highly respected American scientific team. They have set the zoological world, and will very shortly set the whole scientific and lay world, ablaze with excitement. After nearly fifty years of legend and mystery, the saga of the Loch Ness ‘Monster’ is about to end with the addition of a remarkable new (or possibly very ancient) species to the world’s animal kingdom” (Witchell 1975, unpaginated author’s preface).

Dinsdale, despite the rift that would then have existed between himself and Scott, announced his great confidence in the photos (Dinsdale 1973), and such was their apparent significance that they were reported not just in the Nature paper, but on the journal’s cover too. Here is another of Williams’s gaffs, since he describes the paper as an “anonymous item” (p. 175). In fact, Scott and Rines are clearly noted as authors in the article’s abstract (Scott & Rines 1975).

The first part of the Scott & Rines (1975) article. Image: Nature Publishing Group.

The first part of the Scott & Rines (1975) article. Image: Nature Publishing Group.

The publication of this paper is definitely one of the weirder decisions ever made by Nature and one that attracted immediate and strong criticism. Importantly, it makes a mockery of the notion, beloved of cryptozoologists, that ‘the establishment’ has forever shunned or deliberately ignored such things as Nessie. Bullshit, dudes; you had a freakin’ paper in Nature.

The Peter Scott books I own. Image: Darren Naish.

The Peter Scott books I own. Image: Darren Naish.

On that note, one thing that should strike you while reading this book is the extremely high number of working scientists, academic institutions and official societies that, at some time or another since the 1930s, were embroiled in the Loch Ness saga. In, again, massive and hilarious contrast to the idea that academics shun or ignore cryptozoological subjects like the Loch Ness Monster, such qualified workers as Richard Harrison and Leo Harrison Matthews (both well known for their work on marine mammals), marine biologist Sir Alister Hardy, primatologist Sir Solly Zuckerman, palaeontologist Alan Charig*, ethologist, artist, author and TV personality Desmond Morris and many others were all involved or solicited opinion at some point. Hardy, incidentally, stated his belief in the monster (Wiliams 2015, p. 94).

* I wish I’d known this when writing a biography on Charig (Moody & Naish 2010).

Robert Rines has been a mildly controversial figure, and there have been occasions in which his credentials and qualifications were called into question. Here are two letters from the pages of  New Scientist , both from 1982 (vol 95, issues 1315 and 1320, respectively). Image:  New Scientist .

Robert Rines has been a mildly controversial figure, and there have been occasions in which his credentials and qualifications were called into question. Here are two letters from the pages of New Scientist, both from 1982 (vol 95, issues 1315 and 1320, respectively). Image: New Scientist.

On science and scientists, and anti-scientific statements. All in all, A Monstrous Commotion is useful in providing a great deal of novel discussion pertaining to the Scott correspondence, so far so good. But the book is somewhat ruined by a soft pro-Nessie stance that shines through in some places, the author’s insinuation being that Nessie is real and deserving of study and that those scientists and commentators who have rejected its existence and failed to take it seriously are the ones in error.

In places, he appears to unquestionably accept a few notions that, while beloved of Nessie supporters, have been so effectively countered that they shouldn’t ever be used as ‘supporting’ arguments ever again. Examples? That “the Monster [has] a pedigree that [goes] back … over 1,300 years” (p. 9) (see Magin 2001), or that coelacanths can be used to support the idea that the fossil record may as well be disregarded (p. 61).

The idea that  Latimeria , the extant coelacanth, provides support for the view that Mesozoic-grade vertebrate taxa might persist to the present without leaving a fossil record is very naive. In case you hadn’t noticed, we’ve now known of Holocene coelacanths for more than 80 years. This model was on display at the Natural History Museum (London, UK) in 2013. Image: Darren Naish.

The idea that Latimeria, the extant coelacanth, provides support for the view that Mesozoic-grade vertebrate taxa might persist to the present without leaving a fossil record is very naive. In case you hadn’t noticed, we’ve now known of Holocene coelacanths for more than 80 years. This model was on display at the Natural History Museum (London, UK) in 2013. Image: Darren Naish.

Williams notes the sometimes irksome statements made by Nessie supporters about scientists and the scientific process. Constance Whyte, describes Williams, might have seen the scientists of the day as “[a]rrogant and tunnel-visioned”, and that “they could not even be bothered to get out of their armchairs and visit Loch Ness to review the evidence for themselves. Of course, the Monster appeared to flout some basic rules of zoology, and pursuing it could be a waste of time. However, rules were made to be broken, and open-mindedness was supposed to be the hallmark of a good scientist” (p. 63). Williams continues: “All the prejudices and inflexibility of the scientific establishment had been neatly summed up by one of the leading biologists of the 1930s, Sir Arthur Keith FRS” who, Williams tells us, spoke “from the ivory tower of the Royal College of Surgeons” and went on to dismiss the beast “as a problem for psychologists, not zoologists” (p. 63). There are, of course, good reasons for thinking that Keith’s idea of a psychological explanation for monsters is a good one, not the opposite. William’s goes on to refer to the “scientific mafia” when describing the scientific response to Whyte’s 1957 book (p. 63).

The Cryptozoologicon Volume I  sometimes mocks the aggressive tone used by some cryptozoologists. At right, part of John Conway’s scene of a Himalayan yeti group. Image: John Conway/  Conway  et al . (2013)  .

The Cryptozoologicon Volume I sometimes mocks the aggressive tone used by some cryptozoologists. At right, part of John Conway’s scene of a Himalayan yeti group. Image: John Conway/Conway et al. (2013).

This sort of wording will be familiar if you’ve read my 2013 book (co-authored with John Conway and C. M. Kosemen) The Cryptozoologicon. Therein we deliberately mocked the vitriolic, vituperative, frothing-at-the-mouth-with-anger tone aimed at sceptics and working scientists by a certain cadre of cryptozoological believers (Conway et al. 2013). Fact is, the history of research on all the subjects beloved of cryptozoologists – Nessie, bigfoot, the yeti – shows that working scientists never shunned, ignored, dismissed or rejected these things but, on the contrary, spent time considering them, writing about them and even investigating them, only to get their fingers burnt when the subjects proved to mostly be a waste of time (cf Regal 2011). Look again at the list of scientists mentioned above: it’s absolutely farcical to state that scientists haven’t been interested, or haven’t bothered to investigate this stuff. To be clear: Williams isn’t guilty of painting science and scientists in this way, but he’s saying that Whyte was.

Your author (on the right) with Nessie. Image: Darren Naish.

Your author (on the right) with Nessie. Image: Darren Naish.

Indeed, this sort of thing – championed and given the thumbs up by at least some cryptozoologists (and their allies, the paranormalists) – only makes its proponents look naïve and clueless. Caricaturing scientists critical of the Loch Ness Monster as a ‘mafia’ implies that they work together as a band when confronted with a problem. The inner workings and politics of science involve very much the opposite, a fact often destructive and detrimental to those involved. Dear cryptozoologists critical of ‘the scientific establishment’ or of scientific sceptics in general: why do you insist on remaining so clueless with respect to what science is and how it works? There isn’t a gang or club of conspiring scientists who elect to take a given stance on a topic, but a community of competing individuals, all of whom are complex human beings.

The idea that Nessie might have retractable snorkels on its head - an odd idea, to be sure - has long been fairly popular in the LNM literature. The ‘gargoyle head’ photo has to be interpreted within this context. Image: Randall & Keane (1978).

The idea that Nessie might have retractable snorkels on its head - an odd idea, to be sure - has long been fairly popular in the LNM literature. The ‘gargoyle head’ photo has to be interpreted within this context. Image: Randall & Keane (1978).

Finally, a complaint I wish to make about A Monstrous Commotion is that it is, in places, oddly deficient in giving credit. I suppose I shouldn’t expect my own critical comments on various of the LNM photos to warrant mention since (outside of 2016/2017’s Hunting Monsters) they were only ever published here at TetZoo, not in print, but the complete and total absence of Ronald Binns – he isn’t mentioned or even cited once – is odd. Suspiciously so, given that a few sections of the book read much as if they took data or conclusions from Binns (1983). The lack of reference to the many discussions which have occurred within the various ‘parish magazines’ of cryptozoology – Fortean Times, Animals & Men, The Cryptozoology Review, Strange Magazine, Fortean Studies and so on – is also a bit odd.

More evidence for the snorkel-headed Nessie meme: this is the brilliant Kaiyodo toy. Mine was specially shipped from Japan, and oh do I love it. Image: Darren Naish.

More evidence for the snorkel-headed Nessie meme: this is the brilliant Kaiyodo toy. Mine was specially shipped from Japan, and oh do I love it. Image: Darren Naish.

A Monstrous Commotion is an entertaining book that I much enjoyed reading. After a slow start that I would have been happy to go without, it tells a fascinating story and does it well. Those seriously interested in the history of research and ideas on lake monsters should definitely read it, and it might even be said to be one of the best and most professional of books on the Loch Ness Monster yet published. However, it is sometimes appears too sympathetic to those who supported the existence of the Loch Ness Monster, doesn’t appropriately cite all relevant sources, and has enough small, technical errors that it shouldn’t be relied on for factual accuracy.

The only versions of Peter Scott’s renditions of the ‘gargoyle head’ illustration I’ve seen online have been tiny and very low-res, so here’s my best effort at reproducing the best version I have to hand (it’s from the 1981 Reader’s Digest book  Into the Unknown ). Image: Peter Scott/Bradbury (1981).

The only versions of Peter Scott’s renditions of the ‘gargoyle head’ illustration I’ve seen online have been tiny and very low-res, so here’s my best effort at reproducing the best version I have to hand (it’s from the 1981 Reader’s Digest book Into the Unknown). Image: Peter Scott/Bradbury (1981).

Williams, G. 2015. A Monstrous Commotion: the Mysteries of Loch Ness. Orion Books, London. pp. 365. ISBN 978-1-4091-5874-5. Softback, refs. Here at amazon. Here at amazon.co.uk.

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!

Nessie and related issues have been covered on TetZoo a fair bit before, though many of the older images now lack ALL of the many images they originally included…

Refs - -

Bradbury, W. 1981. Into the Unknown. Reader’s Digest Association, Pleasantville, New York/Montreal.

Binns, R. 1983. The Loch Ness Mystery Solved. Open Books, London.

Binns, R. 2017. The Loch Ness Mystery Reloaded. Zoilus Press.

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

Dinsdale, T. 1973. The Rines/Egerton picture. The Photographic Journal April 1973, 162-165.

Dinsdale, T. 1976. Loch Ness Monster, Revised Edition. Routledge & Kegan Paul, London.

Magin, U. 2001. Waves without wind and a floating island – historical accounts of the Loch Ness monster. In Simmons, I. & Quin, M. (eds) Fortean Studies Volume 7. John Brown Publishing (London), pp. 95-115.

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

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

Randall, N. & Keane, G. 1978. Focus on Fact. No. 5 Unsolved Mysteries. W. H. Allen & Co, London.

Regal, B. 2011. Searching for Sasquatch: Crackpots, Eggheads, and Cryptozoology. Palgrave Macmillan, New York.

Rines, R. H. 1982. Summarizing a decade of underwater studies at Loch Ness. Cryptozoology 1, 24-32.

Rines, R. H., Edgerton, H. E., Wyckoff, C. W. & Klein, M. 1976. Search for the Loch Ness Monster. Technology Review March/April 1976, 25-40.

Scott, P. 1976. Why I believe in the Loch Ness Monster. Wildlife 18, 110-111.

Scott, P. & Rines, R. 1975. Naming the Loch Ness monster. Nature 258, 466-468.

Sitwell, N. 1976. The Loch Ness Monster evidence. Wildlife 18, 102-109.

Taylor, M. A. & Martin, J. G. 1990. Big Mouths and Long Necks. Leicestershire Museums, Arts and Records Service, Leicester.

Williams, G. 2015. A Monstrous Commotion: the Mysteries of Loch Ness. Orion Books, London.

Witchell, N. 1975. The Loch Ness Story. Penguins Books, Harmondsworth, Middlesex.

The Cautious Climber Hypothesis

Those of you familiar with the literature on hominid evolution will doubtless have read at least something about the evolution of hominid bipedality. In the most popular model, bipedal hominids originated from terrestrial, chimp-like quadrupeds (which were still capable of climbing but not highly specialised for it), sometime within the last 7 or so million years. However, committed adaptation to full-time bipedality did not occur until more recently, and at least some of the hominids included within the ‘australopithecine’ grade of our lineage (and it obviously is a grade, even Australopithecus itself – as currently conceived – being paraphyletic) were seemingly not far from chimps and bonobos in climbing ability.

Hominids - represented here by a gorilla, orangutan and chimpanzee (the human needed to complete the scene is missing) - are different from other anthropoid primates in many important aspects. What particular adaptational history caused them … us… to be so different? This mural is on show at Edinburgh Zoo, Scotland, and is by Russell Dempster. Image: Darren Naish.

Hominids - represented here by a gorilla, orangutan and chimpanzee (the human needed to complete the scene is missing) - are different from other anthropoid primates in many important aspects. What particular adaptational history caused them … us… to be so different? This mural is on show at Edinburgh Zoo, Scotland, and is by Russell Dempster. Image: Darren Naish.

This shift likely occurred in open habitats, and quite why bipedal adaptation evolved has been the subject of copious speculation. Maybe it was to do with being able to see further, to free up the hands for carrying things, to improve social and/or sexual communication, to assist with thermoregulation, to increase foraging reach, to improve wading abilities or… insert favoured model for origin of bipedality here.

Primate evolution: another of those subjects that gets written about  a lot . Here are some of the (mostly hominid-themed) primate books in the TetZoo collection, but far from all of them. Pet and fringe theories abound in the popular and semi-popular literature on hominid evolution. Image: Darren Naish.

Primate evolution: another of those subjects that gets written about a lot. Here are some of the (mostly hominid-themed) primate books in the TetZoo collection, but far from all of them. Pet and fringe theories abound in the popular and semi-popular literature on hominid evolution. Image: Darren Naish.

What I’ve just described might be regarded as ‘the textbook view’, however, since there are indications that things might not have happened as described. Anatomical details suggest that proficient bipedal abilities might not have originated in open, terrestrial environments, but in wooded habitats, earlier in history, and among taxa that spent more time in trees than on the ground.

There’s a lot more that could be said about that particular area, but here I want to concentrate on an even earlier phase of evolution. Namely, that part relevant to those hominoids in existence prior to the split between pongines (orangutans and their relatives) and hominines (African great apes). Such animals can be termed stem-hominids, pre-hominids or early hominoids, depending on your preference, and they’d be closer in position to hominids than are gibbons (aka hylobatids). I’m going to call them ‘pre-hominids’ since I find this to be the least ambiguous term. Which behavioural, locomotory and ecological specialisations led to the evolution of the hominid body form, and what were pre-hominids like?

The red box shows the section of the family tree we’re especially interested in here. The animals concerned are hominoids, but not part of the hominid crown (that is, they’re not part of the hominid group delimited by living hominid lineages). They’re stem-hominids, or ‘pre-hominids’. Image: Darren Naish.

The red box shows the section of the family tree we’re especially interested in here. The animals concerned are hominoids, but not part of the hominid crown (that is, they’re not part of the hominid group delimited by living hominid lineages). They’re stem-hominids, or ‘pre-hominids’. Image: Darren Naish.

Some authors have proposed that pre-hominids were gibbon-like brachiators (perhaps pre-adapted for bipedality). This is the so-called brachiation, brachionationist or hylobatian model (e.g., Morton 1926, Keith 1934, Tuttle 1981). Others have argued that pre-hominids were more terrestrial, chimp-like knuckle-walkers (e.g., Keith 1923, Washburn 1963), a model similar to the ‘trogolodytian’ one actually proposed as a phase within the hylobatian model. Neither of these models, however, appears consistent with the long list of features that hominids share with mammals that are neither brachiators nor terrestrial quadrupeds, but vertical climbers that use what’s termed an antipronograde posture (where the body’s long axis is oriented more than 45° from the horizontal). The proportionally long arms and short legs of hominids (we’re referring here to the assumed ancestral condition, not that possessed by unusual forms like humans), the short thorax, reduced lumbar region where vertebrae are incorporated into the sacrum, wide shoulders and hips, and the anatomy of the shoulders, wrists and feet all parallel features seen in lorises and kin, and in climbing South American monkeys like spider monkeys (Sarmiento 1995). In short, these features – and there’s other evidence too, relating to the way muscles function and so on – suggest that pre-hominids were perhaps specialised for vertical climbing.

Which form of locomotion was typical of those hominoids ancestral to hominids? Were they brachiators, arboreal climbers, or digitigrade or knuckle-walking terrestrial forms? Image: Richmond  et al . (2001).

Which form of locomotion was typical of those hominoids ancestral to hominids? Were they brachiators, arboreal climbers, or digitigrade or knuckle-walking terrestrial forms? Image: Richmond et al. (2001).

We can go further, since hominids don’t merely possess general features associated with vertical climbing: they have additional features specific to what are known as cautious climbers. These are those vertical climbers that rely on the grasping of (sometimes discontinuously sized) supports and do not leap. Cautious climbers among mammals include lorises, some colobine monkeys, tree sloths and the extinct palaeopropithecid lemurs (Sarmiento 1995) (though some of these taxa – sloths in particular – have specialised for suspensory climbing too). The features we’re talking about here include dorsal migration of the scapula relative to its position on the ribcage in other primates, increased breadth of the manubrium (the big, anterior-most section of the sternum), a reduced number of tracheal rings, a flat-topped diaphragm with a central tendon, a tendon that connects the protective membrane around the heart to the diaphragm (termed the pericardiophrenic tendon), and reduction or loss of the tail.

Did hominids start their history as ‘cautious climbers’, convergently similar to such arboreal mammals as lorises, sloths, various leaf-eating Old and New World monkeys, and some extinct lemurs? The pre-hominid at far right is a hypothetical animal - a ‘concestor’ - that matches this prediction. A slow loris and three-toed sloth are shown at left. Image: Darren Naish.

Did hominids start their history as ‘cautious climbers’, convergently similar to such arboreal mammals as lorises, sloths, various leaf-eating Old and New World monkeys, and some extinct lemurs? The pre-hominid at far right is a hypothetical animal - a ‘concestor’ - that matches this prediction. A slow loris and three-toed sloth are shown at left. Image: Darren Naish.

Because living cautious climbers share a set of physiological, anatomical and behavioural features, we can infer that they were likely present in cautious climbing pre-hominids too. Cautious climbers use slow, deliberate movements, often hold the foot in a hooked pose, use the foot from the toes to the heel when grasping, and are often bipedal when on the ground. Their slow, deliberate and often powerful movements are in keeping with a high proportion of slow twitch or red muscle fibres (Sarmiento 1995).

Cautious climbers also tend to be – but are not always – folivores (leaf eaters), and their reliance on leaves means that they’ve evolved large guts and have slow metabolisms. They’re therefore mostly big (c 10-40 kg), bulky animals with long gestation periods and a reliance on tropical habitats with a guaranteed supply of canopy foliage. Their slow metabolisms also predict that they’re relatively good at dealing with toxins, and it’s been argued that their slow-moving, low-energy strategy makes them inclined to evolve laryngeal specialisations and an ability to broadcast loud sounds over distance.

Cautious climbers generally don’t leap, or drop from height onto other structures when climbing. Instead, they mostly climb slowly and deliberately, with actions like these (here depicted in a potto) being used to move from one branch to another. Image:   Napier & Napier (1985)   (and based on an original by Charles-Dominique).

Cautious climbers generally don’t leap, or drop from height onto other structures when climbing. Instead, they mostly climb slowly and deliberately, with actions like these (here depicted in a potto) being used to move from one branch to another. Image: Napier & Napier (1985) (and based on an original by Charles-Dominique).

The idea that pre-hominids were cautious climbers of this sort, and that the biological and ecological correlates of this adaptive regime were present in these animals, was explored in depth by Esteban Sarmiento in a few papers from the 1990s (Sarmiento 1995, 1998). The significance of the cautious climber model is not just that it better allows us to imagine what pre-hominids might have been like; it’s also interesting in that imaging hominids as animals that went through this phase helps explains why they were evolutionary shaped in the ways that they were.

Hominid skeletons - this is that of an orangutan, photographed at Musée d'histoire Naturelle, Tournai (Belgium) - possess numerous features indicative of arboreal adaptation. Image: Michel Wal, CC BY-SA 3.0, wikipedia (original  here ).

Hominid skeletons - this is that of an orangutan, photographed at Musée d'histoire Naturelle, Tournai (Belgium) - possess numerous features indicative of arboreal adaptation. Image: Michel Wal, CC BY-SA 3.0, wikipedia (original here).

Hominids took to frugivory on later occasions within their history, for example, but a folivorous initial phase might explain why they have the teeth that they do (the incisors are relatively small, the canines are shortened, there are crushing surfaces on the premolars, enamel wrinkling is present on the molars, and so on), why the front of the face is reduced, and why the lower jaw has such a large and vertical ascending process, a big section adjacent to the molars and a broad but short condyle (Sarmiento 1995). A folivorous heritage could also help explain why hominids are relatively slow growing, resistant to many poisons toxic to other mammals, and equipped with an ability to make loud, complex calls. It would appear, however, that pre-hominids were not as specialised for folivory as are cautious climbers like sloths, since hominids never evolved a complex stomach and always maintained good terrestrial abilities.

Was Sarmiento right about cautious climbing and folivory being all that important in primates close to the ancestry of later hominid lineages? This isn’t an area that’s been all that intensively discussed but – when comments on ‘pre-hominid’ evolution have been provided – authors have tended not to state special preference for the idea, at least considering it as likely as suspensory behaviour or generalised climbing (e.g., Pilbeam & Young 2004, Begun 2016). Richmond et al. (2001), however, noted that ideas positing antipronograde climbing postures were “[a]rguably the most popular [of evolutionary models pertaining to pre-hominid lifestyle] … during the last several decades” (p. 81) and argued that the anatomy and biomechanics of extant hominids are consistent with an arboreal climbing ancestry. They didn’t specifically have cautious climbing and/or folivory in mind though.

This article isn’t about SpecBio. But if pre-hominids were vaguely sloth-like in some ways, a different trajectory of hominid evolution might have resulted in a radiation of increasingly sloth-like primates… in which case, maybe things like this could have evolved. Maybe. This is  Giganthropus , a fictional sloth-like hominid featured in   Dougal Dixon’s 1990  Man After Man   . Image: Philip Hood, in   Dixon (1990)  .

This article isn’t about SpecBio. But if pre-hominids were vaguely sloth-like in some ways, a different trajectory of hominid evolution might have resulted in a radiation of increasingly sloth-like primates… in which case, maybe things like this could have evolved. Maybe. This is Giganthropus, a fictional sloth-like hominid featured in Dougal Dixon’s 1990 Man After Man. Image: Philip Hood, in Dixon (1990).

As goes fossils, animals that pertain to the right approximate part of the cladogram do seem to provide support for the model. Morotopithecus from Early Miocene Uganda is large (20-40 kg) (Gebo et al. 1997) and has been inferred to have been a vertical climber (and perhaps a cautious climber), while Pierolapithecus from the middle Miocene of Spain has a set of features which appear inconsistent with suspensory behaviour and more in line with vertical climbing (Moyà-Solà et al. 2004) (and, again, perhaps with cautious climbing too).

All in all, I find the ‘cautious climber’ model of pre-hominid evolution intriguing and worthy of more attention, hence this article. Long-term readers might remember me mentioning this article – as an in-prep piece, waiting completion – for some years now. It is, at last, ticked off the list.

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

For previous TetZoo articles on primate diversity and evolution, see…

 Refs - -

Begun, D. R. 2016. The Real Planet of the Apes: a New Story of Human Origins. Princeton University Press, Princeton and Oxford.

Dixon, D. 1990. Man After Man: An Anthropology of the Future. Blandford, London.

Gebo, D. L., MacLatchy, L., Kityo, R., Deino, A., Kingston, J. & Pilbeam, D. 1997. A hominoid genus from the Early Miocene of Uganda. Science 276, 401-404.

Keith, A. 1923. Man’s posture: its evolution and disorders. British Medical Journal 1, 451-454, 499-502, 545-548, 587-590, 624-626, 669-672.

Keith, A. 1934. The Construction of Man’s Family Tree. Watts, London.

Morton, D. J. 1926. Evolution of man’s erect posture: preliminary report. Journal of Morphology and Physiology 43, 147-149.

Moyà-Solà, S., Köhler, M., Alba, D. M., Casanovas-Vilar, I. & Galindo, J. 2004. Pierolapithecus catalaunicus, a new Middle Miocene great ape from Spain. Science 306, 1339-1344.

Napier, J. R. & Napier, P. H. 1985. The Natural History of the Primates. British Museum (Natural History), London.

Pilbeam, D.  & Young, N. 2004. Hominoid evolution: synthesizing disparate data. C. R. Palevol 3, 305-321.

Richmond, B. G., Begun, D. R. & Strait, D. S. 2001. Origin of human bipedalism: the knuckle-walking hypothesis revisited. Yearbook of Physical Anthropology 44, 70-105.

Sarmiento, E. E. 1995. Cautious climbers and folivory: a model of hominoid differentiation. Human Evolution 10, 289-321.

Sarmiento, E. E. 1998. Generalized quadrupeds, committed bipeds, and the shift to open habitats: an evolutionary model of hominid divergence. American Museum Novitates 3250, 1-78.

Tuttle, R. H. 1981. Evolution of hominid bipedalism and prehensile capabilities. Philosophical Transactions of the Royal Society 292, 89-94.

Washburn, S. L. 1963. Behavior and human evolution. In Washburn, S. L. (ed) Classification and Human Evolution. Aldine, Chicago, pp. 190-201.

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

Listeners of the Tetrapod Zoology podcast will know that I have – for no specific reason – been going through a bit of a Loch Ness Monster phase recently, my stated aim being to review three recently-ish published books on the subject. Here’s the first of those reviews, devoted to Ronald Binns’s The Loch Ness Mystery Reloaded. I provided a very brief review of this book at TetZoo ver 3 back in 2017 but always planned to produce a longer version. So here we are.

Nessie, the beast of many forms. It’s almost as if people are describing all manner of different things. Images: Darren Naish.

Nessie, the beast of many forms. It’s almost as if people are describing all manner of different things. Images: Darren Naish.

Is the Loch Ness Monster (LNM) science? Should I be writing about it, and encouraging an interest in it, on a blog ostensibly devoted to the scientific study of the natural world? Here I’ll say what I’ve said several times before about monsters and cryptozoology in general: even if monsters don’t exist (in the zoological sense), there’s still a phenomenon here that’s worthy of study, and there’s still a body of data that we can subject to scientific analysis. And I’ll add that ideas and writings about monsters like Nessie are definitely relevant to those of us intrigued by speculative zoology (Naish 2014), fringe theories, the history of zoology and other subjects included within the TetZoo remit.

Binns (1983)  , hardback (at left) and 1984 softback edition.

Binns (1983), hardback (at left) and 1984 softback edition.

Ronald Binns’s 1983 The Loch Ness Mystery Solved – produced with assistance from Rod Bell (though he doesn’t get an authorship credit) – is a classic work of scholarship and scepticism (Binns 1983). It shows how sightings, photos and film purporting to describe or show the monster are less impressive than typically described, are indeterminate or of more prosaic identity than claimed, have been embellished or modified by enthusiastic or biased writers, and can sometimes be explained as encounters with known animals (seals, waterbirds, deer). A sociocultural setting for the monster, an evaluation of the clutching-at-straws ideas on its biology, appearance, phylogenetic affinities and ecology, and a takedown of the ‘historical Nessie’ endorsed elsewhere also feature within the book (Binns 1983).

It should not be assumed that people - even people who’ve lived their lives in rural places, surrounded by wildlife - can always identify such animals as deer, seals and waterbirds (like grebes and cormorants) when they see them in unusual places, poses or situations. Deer are abundant around Loch Ness. I photographed this male Red deer adjacent to Loch Knockie, which is just a few hundred metres to the east of Loch Ness. Image: Darren Naish.

It should not be assumed that people - even people who’ve lived their lives in rural places, surrounded by wildlife - can always identify such animals as deer, seals and waterbirds (like grebes and cormorants) when they see them in unusual places, poses or situations. Deer are abundant around Loch Ness. I photographed this male Red deer adjacent to Loch Knockie, which is just a few hundred metres to the east of Loch Ness. Image: Darren Naish.

I’m glad to have encountered it at a relatively early phase in my career as a monster researcher, since virtually everything else I read and was exposed to was extremely pro-monster, so much so that I spent time as a teenager thinking that Nessie was a scientific likelihood. Phew: lucky, then, that I never made a fool of myself by proclaiming a belief in Nessie. Oh, wait.

Of course, much has happened since the publication of Solved in 1983. We have new biographic information on the people integral to the LNM story, and new eyewitness accounts, illustrations and photos have been unearthed or have joined the collective pool. Several of the main characters in the LNM story have died since Solved was published, meaning that their role can be more fully and honestly assessed than when they were alive (Binns 2017). On the sociological angle, Nessie has remained a cultural icon and flashpoint for woo, and its story has been retold, embellished, added to and expanded via the publication of many post-1983 books, so many that there’s what looks like a (mostly British) cottage industry on the subject.

Cover of   Binns (2017)  . Buy it if interested in the Loch Ness Monster, lake monster lore, cryptozoology or scepticism.

Cover of Binns (2017). Buy it if interested in the Loch Ness Monster, lake monster lore, cryptozoology or scepticism.

In view of all this, the time is right for an addendum to Solved, and thus we find The Loch Ness Mystery Reloaded (Binns 2017). Reloaded is, to quote its author, essentially a long-form appendix to The Loch Ness Mystery Solved, but it isn’t at all dry or tedious. It’s well written, entertaining and absorbing if you’re familiar with any aspects of LNM lore, and in fact is probably the LNM-themed book I’ve enjoyed reading the most. Footnotes and references are provided, though there are only a handful of illustrations.

The book begins with a look back at the making of Solved, the responses to it, and a series of updates on the book’s main (human) characters. Binns then retreads two classic sightings – that of Donaldina Mackay and the Spicers – albeit with new information. The Spicer account of 1933 is the great classic ‘land sighting’, recounted in every Nessie-themed book. George Spicer is generally framed as a lucky everyman who saw a remarkable, inexplicable thing and reported it fairly, without fanfare and without any need to see himself – rather than the animal – as the point of interest.

The Spicer sighting as conventionally portrayed, here by Dinsdale (1976). Image:   Dinsdale (1976)  .

The Spicer sighting as conventionally portrayed, here by Dinsdale (1976). Image: Dinsdale (1976).

None of this is correct. Details of Spicer’s account make it likely that he and his (still nameless) wife saw bounding deer (an idea, not original to me, discussed in Hunting Monsters); the details of his sighting changed significantly over the years and demonstrate (I say again: demonstrate) both embellishment and a predilection on Spicer’s part to speculate. Furthermore, Spicer wasn’t a quiet, impartial witness, reluctantly discussing his encounter when door-stepped by thirsty journalists, but extraordinarily enthusiastic, garrulous and boastful about it, his own writings making him sound wide-eyed and credulous. There’s a popular idea among cryptozoologists that one should ignore personality traits and biography and just pay attention to the monster sighting. Alas, no; this is wrong. Those things are absolutely relevant.

Probably the best ever depiction of the Loch Ness Monster in action. Surely this is what Mr Spicer actually saw. A beautiful image by the legendary Gino D’Achille. Image: (c) Gino D’Achille.

Probably the best ever depiction of the Loch Ness Monster in action. Surely this is what Mr Spicer actually saw. A beautiful image by the legendary Gino D’Achille. Image: (c) Gino D’Achille.

Famous photos, reloaded. Chapters are also devoted to the more notable LNM photos and the Dinsdale film. Revised and updated takes on the Surgeon’s photo, and on the Gray, Stuart, Cockrell, Macnab and O’Connor photos – and a bunch of less famous ones – are presented, all of which can sensibly be stated to be hoaxes, indeterminate, waves, sticks and other non-animals. Binns remains non-committal on the very unusual Peter O’Connor photo of 1960 but implies that O’Connor’s career as a taxidermist might mean that we’re looking at a dead animal (maybe a seal?). My competing idea (Naish 2017) – it originates with Dick Raynor – is that it’s an inverted kayak and a head-shaped stick in front. I think this explains what looks like planking and the metal rudder support, visible in uncropped versions of the image.

The O’Connor photo has never made sense as goes lighting. O’Connor’s story is suspiciously odd. And the object he photographed does not appear to be an animal. It looks likely to be an inverted kayak, and looks uncannily similar to the specific kayak that O’Connor owned. The image here - scanned from one of the several LNM books - is a cropped version that doesn’t feature the whole of the object. Image: (c) Peter O’Connor.

The O’Connor photo has never made sense as goes lighting. O’Connor’s story is suspiciously odd. And the object he photographed does not appear to be an animal. It looks likely to be an inverted kayak, and looks uncannily similar to the specific kayak that O’Connor owned. The image here - scanned from one of the several LNM books - is a cropped version that doesn’t feature the whole of the object. Image: (c) Peter O’Connor.

Nor does Binns wholly buy my argument in Hunting Monsters than the Hugh Gray photo depicts a swan. He does think it’s a fake though. I remain pretty confident about the swan hypothesis because a Whooper swan (not a Mute swan as implied by the illustration in Hunting Monsters) matches the shape, hue and likely size of the object, the object has what looks like a swan’s ankle sticking out of its flank in the right position, and also has an otherwise inexplicable pale patch (presumably double-exposed on top of the main image, and coming from a moment in time when the bird had its head above the water surface) that exactly matches a Whooper swan’s bill patch. It also has a swan’s tail. And it’s white.

At top, the Hugh Gray photo of 1933. A ‘mid-sized’ object (look at the ripples), that’s white or near-white, has a sinuous appendage at one end, a short, pointed appendage at the other, and a dark appendage that disappears into the water close to one of its ends. The double-strike areas (featuring pale triangular patches) indicate double-exposure (that is, the film failed to move on and was exposed again). Image:   Naish (2017)  .

At top, the Hugh Gray photo of 1933. A ‘mid-sized’ object (look at the ripples), that’s white or near-white, has a sinuous appendage at one end, a short, pointed appendage at the other, and a dark appendage that disappears into the water close to one of its ends. The double-strike areas (featuring pale triangular patches) indicate double-exposure (that is, the film failed to move on and was exposed again). Image: Naish (2017).

On vocal champions. It’s frequently implied or stated by those endorsing the existence of monsters that vocal champions of the cause are sensible, level-headed, scholarly types who come to the conclusions they do after carefully and scientifically evaluating the data. Maybe this is true for some individuals. But it’s generally the opposite of true. Monster proponents – and I’ve now been bold enough to state or imply this in print (Conway et al. 2013, Naish 2017) – are more frequently tremendously naïve, biased, impressionable and unscientific, and attracted to the subject because of its intrinsic appeal and their own tendency to want the world to be occupied by stuff considered beyond current scientific knowledge.

This is emphasised in Binns’s chapters on F. W. ‘Ted’ Holiday, author of 1968’s The Great Orm of Loch Ness (wherein Nessie is posited to be a giant Tullimonstrum) and 1973’s The Dragon and the Disc (the main thesis of which links lake monsters with the ancient alien movement). Similarly, Binns’s treatment of Tim Dinsdale, the well-meaning and likeable champion of the LNM from 1960 until his death in 1987, discusses Dinsdale’s propensity to be all too impressionable, and with ideas that were not in keeping with his claim that he was always objective and basing everything monster-related on sound science. There’s a lot that could be said about both men (I’m avoiding that here; Dinsdale will be discussed in a future article); anyone interested in their legacy and how it fits into the LNM story must see what Binns says to say.

Ahh, classic Holiday. The object in the Hugh Gray photo interpreted as a modern-day, big  Tullimonstrum . Image:   Holiday (1968)  .

Ahh, classic Holiday. The object in the Hugh Gray photo interpreted as a modern-day, big Tullimonstrum. Image: Holiday (1968).

Both those interested in, and brought to despair by, the literature on the LNM will have noted that there’s a lot of it: an enormous number of published books, booklets and articles, and a vast quantity of correspondence, imagery and art. Yet for all the fame of the LNM, there’s been surprisingly little effort to collate or gather things and several collections are in danger of being lost when their owners are no more and their estates dissolved. Binns terms this the ‘archive problem’, and it’s common to many subject areas that involve ‘paperwork’.

But a fraction of the books that exist on the Loch Ness Monster. Image: Darren Naish.

But a fraction of the books that exist on the Loch Ness Monster. Image: Darren Naish.

As mentioned above, a great many books have been published on the LNM since the early 1980s. Binns provides commentary on the more recent of them (those from the post-Dinsdale era). He's fairly critical of Gareth Williams’s serious and well-drafted A Monstrous Commotion of 2015 (another of the books set to be reviewed here), noting that it fails to credit The Loch Ness Mystery Solved despite drawing substantially upon its contents. Those lines in the acknowledgements of A Monstrous Commotion explaining how eagle-eyed reviewers and editors did a sterling job in preventing numerous mistakes are ironic in view of the fairly enormous number of errors that did make it into print, all of which Binns discusses and corrects (pp. 150-152).

Binns on Hunting Monsters. I am of course especially interested (I hope understandably) in what Binns says about my own Hunting Monsters, a volume he classes as part of the same ‘cultural cryptozoology’ school as Loxton and Prothero’s Abominable Science! (which I had a hand in as technical reviewer and blurb-writer; my review of that book is here at TetZoo ver 3). Hunting Monsters is “fluent, elegant, scholarly”, says Binns (p. 153); he “finds Naish’s approach congenial because he seeks to rescue cryptozoology from the naïve literalists” (p. 154).

In which I tried to develop a sociological or anthropological view of cryptids - Nessie included - as icons embedded within culture. The softback version was preceded by a 2016 digital one (which has a different cover). Image:   Naish (2017)  .

In which I tried to develop a sociological or anthropological view of cryptids - Nessie included - as icons embedded within culture. The softback version was preceded by a 2016 digital one (which has a different cover). Image: Naish (2017).

It isn’t a coincidence that Binns finds my take on monsters concordant with his own position. As has been stated here at TetZoo many times now, I think today that cryptids like Nessie are sociocultural phenomena, that people ‘see’ monsters like Nessie because they fully expect to encounter a thing that’s firmly embedded within their culture (Binns terms this expectant attention), that ideas about the identity, biology, ecology and so on of Nessie are nothing more than speculative house-of-cards-type efforts, and that eyewitness descriptions of Nessie are either so ambiguous as to be useless, are hoaxes, or are explainable or near-explainable by those able to employ critical thinking or sceptical analysis.

You can be an atheist and still love buildings of worship. Likewise, thinking that Nessie is not real does not stop Loch Ness from being a remarkable place or one with a great amount of allure. Image: Darren Naish.

You can be an atheist and still love buildings of worship. Likewise, thinking that Nessie is not real does not stop Loch Ness from being a remarkable place or one with a great amount of allure. Image: Darren Naish.

The Age of the Internet. It’s no secret among those especially interested in the LNM that one of the most visible writers on the subject in the Age of the Internet is not a journalist or scientist but a blogger – Roland Watson – who is unashamed and thoroughly biased in his insistence that Nessie is real and that those questioning the ‘evidence’ are wrong and on ground shakier than the firm footing occupied by himself. Binns notes that Watson has been useful in uncovering new data but more impressive are the substantial number of occasions in which Binns corrects or contests Watson’s interpretations.

Watson’s blog is a fun read if you like being immersed in detailed discussions of LNM-themed sightings, anecdotes, accounts and controversies. I congratulate anyone aiming to put esoteric data like this on the record, and to unearth more information on relevant people and the backgrounds to claimed eyewitness accounts.

But two issues ruin Watson’s entire take on the subject. One is his obvious employment of confirmation bias and insistence that Occam’s Razor has no place in monster hunting: while the record shows that he doesn’t automatically identify any object seen in the waters of Loch Ness as a monster, his primary argument on many occasions is to insist that ‘giant unknown aquatic vertebrate species’ should, as an explanation, be on equal footing with ‘swimming deer’, ‘unidentified fish’ or ‘wave’. No it shouldn’t.

The surface of Loch Ness, photographed in September 2016. Many people’s monster sightings are of a calibre similar to this. These are waves, made by a boat. Image: Darren Naish.

The surface of Loch Ness, photographed in September 2016. Many people’s monster sightings are of a calibre similar to this. These are waves, made by a boat. Image: Darren Naish.

The second issue is Watson’s obsession with sceptics, who he very much regards as The Bad Guys, The Enemy. He bashes sceptics a lot – as if being sceptical about Nessie is a bad thing or represents poor life choice – and has written at length about the motivations that, he thinks, drive sceptics and their scepticism. These arguments are naïve, way off base and just fucking weird. Example: Watson claimed on his blog and in his review of Hunting Monsters that I have been groomed – his word – by Loch Ness expert Dick Raynor, since sceptics require that the chalice of Loch Ness scepticism be passed down across the generations, and, furthermore, that I have been turned to the cult of scepticism via the nefarious chicanery and allure of an older man. Binns noted all of this as it was happening. “Naish’s assertion that Watson inhabits ‘an idiosyncratic intellectual landscape’ was a generous and measured response”, he says (p. 18). Damn straight.

Those of us sceptical about lake monsters are so, or have become so, because we aren’t convinced by the evidence, such as it is, and we see obvious problems with the ‘data’ deemed integral to the case by believers. I’ve become a sceptic for these reasons, not because I wish to be part of a special club (note to Nessie fans reading this: I am not employed in academia), not because I somehow earn points or money for being a sceptic (I wish), and not because I’m guilty of sloppy thinking, blindness, unfamiliarity with the evidence or an unwillingness to read or engage with the literature or the cryptozoological community. I was a believer once, I remind you, and I’ve said all sorts of dumb things supporting the existence of monsters that my critics never seem to be aware of. They only see an enemy: a blinkered ivory-tower elite who lives in a solid gold house, has never had a proper job, and dives every evening into a money pit of Science Dollars provided by the government or something.

Here’s a close-up of the Hugh Gray photo again. I say again: the ripples show that this object is ‘mid-sized’ (as in, 1 m long or so). Weird than an object which I say looks like a swan also has what looks like a swan’s leg and what looks like a swan’s pointed tail.

Here’s a close-up of the Hugh Gray photo again. I say again: the ripples show that this object is ‘mid-sized’ (as in, 1 m long or so). Weird than an object which I say looks like a swan also has what looks like a swan’s leg and what looks like a swan’s pointed tail.

You should buy this book. The Loch Ness Mystery Reloaded is required reading for those interested in the Loch Ness phenomenon, in lake monsters, cryptozoology and/or scepticism, and I really enjoyed it. My one complaint is that the lack of an index is a real pain. We all know that indexes are difficult and time consuming to compile, but lacking one entirely is not really excusable – even a short one consisting of keywords is better than nothing.

Anyway… it would be helpful to read Reloaded right after The Loch Ness Mystery Solved if you haven’t read that book already, but I wouldn’t say that this is a requirement. You should definitely buy it. And you should also consider leaving a positive amazon review given a really appalling bias present there at the time of writing. More Nessie book reviews soon.

Binns, R. 2017. The Loch Ness Mystery Reloaded. Zoilus Press. pp. 222. ISBN 978-1-9997359-0-6. Softback, refs. Here at amazon, here at amazon.co.uk.

Nessie and related issues have been covered on TetZoo a fair bit before, though many of the older images now lack ALL of the many images they originated included…

Refs - -

Binns, R. 1983. The Loch Ness Mystery Solved. Open Books, London.

Binns, R. 2017. The Loch Ness Mystery Reloaded. Zoilus Press.

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

Dinsdale, T. 1976. Loch Ness Monster (Revised Edition). Routledge & Kegan Paul, London.

Holiday, F. W. 1968. The Great Orm of Loch Ness. Faber and Faber, London.

Loxton, D. & Prothero, D. R. 2013. Abominable Science! Columbia University Press, New York.

Naish, D. 2014. Speculative zoology. Fortean Times 316, 52-53.

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

Potoos As An Internet Phenomenon

This might be an unusual TetZoo article. I like memes, by which I mean the internet images or concepts that are passed ‘virally’ from one person to another and become modified and/or added to across their ‘generations’. As you’ll know if you pay any attention to memes (or are part of any community where memes are frequently encountered), several animal species are meme favourites. Among them are potoos.

It’s true, I’m guilty of using potoos in memes myself. The one at left was my profile picture on social media for a while; the one at right appeared at TetZoo ver 3. The original photo has been memed a lot. The original photo is sometimes credited to Kristin Lundquist, though that’s not where I got it from. Image: (c)  Kristin Lundquist .

It’s true, I’m guilty of using potoos in memes myself. The one at left was my profile picture on social media for a while; the one at right appeared at TetZoo ver 3. The original photo has been memed a lot. The original photo is sometimes credited to Kristin Lundquist, though that’s not where I got it from. Image: (c) Kristin Lundquist.

Potoos – officially nyctibiids, and not to be confused with the primates called pottos – are a South, Central and North American group of cryptically coloured, nocturnal, big-eyed, insectivorous birds. Nearly all of the seven extant species* are included within the same genus (Nyctibius), the exception being the Rufous potoo N. bracteatus which was given its own genus (Phyllaemulor) in 2017. In addition, there are fossil potoos (from Europe and North America) which have been given separate genera too. Potoos hunt by aerial hawking or by flycatching from a perch, their prey mostly involving beetles and moths. They’ve also been reported to eat small passerines: a Northern potoo N. jamaicensis was once found to contain a White-collared seedeater Sporophila toqueola in its stomach (Cleere & Nurney 1998). But they’re also famously sedentary, spending the day sitting still on a branch or stump where their plumage, shape and posture helps conceal them from predators. Unique notches in their eyelids enable them to see even when the eyelids are closed, and their short, stocky tarsometatarsi appears to be an adaptation for long periods of sitting still.

* Genetic variation within various of these species suggests that this count is too low and it’s very likely that a set of cryptic species are awaiting recognition (Cohn-Haft 1999).

Potoos have an obvious similarity with nightjars, frogmouths and kin, and hence have traditionally been classified alongside them within Caprimulgiformes. While there’s an awful lot to say about caprimulgiform evolution and diversity and where these birds fit within the neornithine family tree, we’ll ignore all that for now since that’s not why we’re here.

Two of a set of famous potoo-themed photos (there are at least four in total). These went viral in 2015 and finding a photographer to credit has not been possible, but if there’s anyone in particular who should be credited, I’d be interested in knowing their name. This is definitely a Great potoo  Nyctibius grandis  (or, part of the series of populations currently included within that species, anyway).

Two of a set of famous potoo-themed photos (there are at least four in total). These went viral in 2015 and finding a photographer to credit has not been possible, but if there’s anyone in particular who should be credited, I’d be interested in knowing their name. This is definitely a Great potoo Nyctibius grandis (or, part of the series of populations currently included within that species, anyway).

Potoos, frankly, sometimes look scary. In the Great potoo N. grandis, the giant eyes have dark brown irides, meaning that they sometimes look entirely black. This is a marked difference from other potoos, all of which have orange, yellowish or yellow-green irides, though the Rufous potoo is also unusual, this time in having a dark vertical wedge in the lower half of its iris (Cohn-Haft 1999). The Great potoo – the largest of the family – is also odd in being big, reaching 54 cm in total length and with a wingspan of around 80 cm. I guess it’s not called the Great potoo for nothing. A bird of this sort, at this size, is somewhat daunting.

Potoos and some amused (or bored) owls. I love this picture. It’s   here on imgur  . Image:  princeemberstorm .

Potoos and some amused (or bored) owls. I love this picture. It’s here on imgur. Image: princeemberstorm.

But potoos can also look comical, cute and more like puppets or cartoon characters than real animals, so – conversely – they also appeal to us for those reasons. Their giant, often bright orange eyes sometimes look like billiard balls and appear unrealistically large within their heads. A small, slim, curved bill accentuates this big-eyed look, though when I talk of the bill here, I’m referring to the strongly hooked anterior portion and not to the wide gape, virtually all of which is concealed when the bird has its mouth closed.

Potoos are just ripe for things like this, it seems. This image, titled ‘Potoo is a sir’, appeared on imgur in 2014 and is   borrowed from here  . Image: (c)  Polarbearsarentreal .

Potoos are just ripe for things like this, it seems. This image, titled ‘Potoo is a sir’, appeared on imgur in 2014 and is borrowed from here. Image: (c) Polarbearsarentreal.

Add all of this together and we come to the reason for this article. Within recent years, potoos have been cartoonified, memed and caricatured to hell, so much so that we might talk of The Potoo As An Internet Phenomenon. I don’t know if any of this happened to potoos before there was an internet, but I don’t see evidence that it did. What’s happened here is the same as that for anything we regard as an internet meme: people saw some cool, interesting pictures and were inspired to depict their own take on them, and other people were inspired by those images, and we then get the domino effect whereby things get mixed and matched, elaborated, or taken in a whole new direction. Keep reading to see where this goes.

Potoo-themed accounts of Twitter. The avatar at left is that of illustrator   Angela Hsieh, aka @hisiheyah  . A selection of others are at right:   our lord and saviour potoo or @PotooPotoo   is the most relevant for the purposes of the article here. Images: Angela Hsieh, Twitter.

Potoo-themed accounts of Twitter. The avatar at left is that of illustrator Angela Hsieh, aka @hisiheyah. A selection of others are at right: our lord and saviour potoo or @PotooPotoo is the most relevant for the purposes of the article here. Images: Angela Hsieh, Twitter.

Part of the story of the potoo’s popularity onlines comes from the appearance of a series of photos, apparently taken in Venezuela in 2013, that show a Great potoo sat on a fence in various poses (two are shown above). The bird looks especially striking in these photos due to its massive and seemingly fully black eyes, its (perhaps surprisingly large) size, and a streamlined look which maybe makes it look somehow unnatural. Plus, it’s a potoo. I mean, seriously, they all look striking if you haven’t seen one before. On size: we can see from accompanying objects (barbed wire, humans hands) that the bird isn’t really that big, but it was big enough (ha ha, another meme) to inspire this, which I like very much…

‘The Great Potoo’, interpreted somewhat literally, from the online comic Question Duck (  original panel here  ). Image: (c)  Question Duck .

‘The Great Potoo’, interpreted somewhat literally, from the online comic Question Duck (original panel here). Image: (c) Question Duck.

The popularity of these photos is also partly explained by the fact that they were used in some bullshit story about a ‘dinosaur owl’ being resurrected from ancient DNA (there’s even a Snopes takedown of that concept), plus people who’ve never heard of books (let alone potoos) have claimed variously that the animal might be an alien, a demon or a piece of created taxidermy. Karl Shuker discussed his investigation of the photos at his blog, here.

Where have people gone from there? Some of you will know that griffons … or griffins, or gryphons, or gryphins … are big on the internet these days. For whatever reason, they’ve brought out people’s creative side, and there are griffin-inspired creatures of all sorts, depicted variously in attractive fantasy art, in pokemon-style cartoonish art, in anatomically rigorous hyperdetailed art, and even in furry costume form. I can’t mention furry costumes without bringing your attention to the fact that, yes, at least one person has made a potoo cosplay, and it’s spectacular. Here’s Casanova the potoo, as made by the fantastic people at Menagerie.

Other potoo-themed costumes are now available - I’m pretty sure that Casanova is not a one-off. Image: (c) SarahDee/Fur Affinity, original   here  .

Other potoo-themed costumes are now available - I’m pretty sure that Casanova is not a one-off. Image: (c) SarahDee/Fur Affinity, original here.

But back to griffins. I say that griffins are interesting “for whatever reason”, but the real reason, of course, is that they’re awesome, combining the most visually compelling parts of some of the world’s most attractive and popular animals, by which I mean big raptors and big cats. I’ve always really liked griffins myself, and I remember being thrilled when discovering that people even had the audacity to invent an even-more-elaborate griffin spin-off, the Hippogriff!

Combine the attractive qualities of griffins with a new Golden Age of inventive creature art, and with the existence of potoos, and with the fact that potoos are an internet phenomenon, and the inevitable consequence is … potoo griffins. Yes, there are many potoo griffins out there. Here are just two illustrations of them…

Cartoon potoo griffins, by Cyndi Foster, aka Gingo, aka Gryphon Queen (  here at DeviantArt  ). This is why the internet was made. Images: (c)  Cyndi Foster/Gingo .

Cartoon potoo griffins, by Cyndi Foster, aka Gingo, aka Gryphon Queen (here at DeviantArt). This is why the internet was made. Images: (c) Cyndi Foster/Gingo.

And this sort of thing – the combining of animal parts to create chimaeras that stun or surprise us by virtue of how absurd, or silly or compelling or realistic or wonderful they are – has led to yet more potoo-inspired creatures of the internet. I give you: a potoo x marbled polecat chimaera, a potoo x black panther chimaera, a potoo x Pallas cat chimaera, and a potoo x Chihuahua chimaera. Brilliant (and all credit to the artists). There are many, many others.

At left, a marbled polecat x potoo griffin, “the most majestic and noble of griffins”, by uropygid (original  here ). At right, a potoo griffin that combines black panther and potoo features, by Seneca’s Art Rocks. Images: (c)  Uncle Max/uropygid ,  Seneca’s Art Rocks .

At left, a marbled polecat x potoo griffin, “the most majestic and noble of griffins”, by uropygid (original here). At right, a potoo griffin that combines black panther and potoo features, by Seneca’s Art Rocks. Images: (c) Uncle Max/uropygid, Seneca’s Art Rocks.

Potoo x Pallas cat by iguanamouth (from their “kept getting requests for gryphons so heres a bunch of them At Once “ project), Potoo x Chihuahua by Pechschwinge, from their Daily Gryphon Challenge. Images: (c)  iguanamouth ,  Pechschwinge .

Potoo x Pallas cat by iguanamouth (from their “kept getting requests for gryphons so heres a bunch of them At Once “ project), Potoo x Chihuahua by Pechschwinge, from their Daily Gryphon Challenge. Images: (c) iguanamouth, Pechschwinge.

 Yes, truly we are in the age of peak potoo, a time when we might pay reverence to Our Lady Potoo of the Sacred Heart, or Our Lord and Savior Potoo Bird, or something along those lines. What’s that, you say – this would work as a great internet image as well? Well…

‘Our Lord and savior potoo bird’. This image is by dragongirl222, who does   a whole range of potoo-themed merchandise at redbubble  . Image: (c)  dragongirl222 .

‘Our Lord and savior potoo bird’. This image is by dragongirl222, who does a whole range of potoo-themed merchandise at redbubble. Image: (c) dragongirl222.

This, of course, is not everything. There’s even more out there — much more — and it’s easy to find if you go look for it. Dear reader: I give you… The Potoo As An Internet Phenomenon.

Thanks to those supporting this work – and the very blog itself – via pledges at patreon. You can support what I do and see works-in-prep behind the scenes, via pledges as small as $1 per month.

For previous TetZoo articles relevant to potoos, see…

Refs - -

Cleere, N. & Nurney, D. 1998. Nightjars: A Guide to Nightjars and Related Nightbirds. Pica Press, Mountfield.

Cohn-Haft, M. 1999. Family Nyctibiidae (potoos). In del Hoyo, J., Elliott, A. & Sargatal, J. (eds) Handbook of the Birds of the World, Vol. 5. Barn-owls to Hummingbirds. Lynx Edicions, Barcelona, pp. 288-301.

Recollections of Dinosaurs Past and Present, the 1980s Exhibition

As a regular reader here, you should be familiar with my interest in the portrayal of dinosaur life appearance, and indeed in palaeoart in general. It’s now trite to explain how the iconic, well-drafted, anatomically rigorous illustrations of Robert Bakker, Greg Paul, Mark Hallett and others were integral to the popularisation and dissemination of the ‘dinosaur renaissance’ that occurred between the late 1960s and 1990s, and few interested in prehistoric life will have failed to notice how quickly and frequently reconstructions of extinct species – often good, accurate and innovative – appear today, typically in the digital medium. Palaeoart remains relevant, essential, and with a huge fanbase.

Robert Bakker’s 1969 sprinting  Deinonychus , produced to accompany John Ostrom’s seminal article on this amazing dinosaur. Artistic depictions like this one cement the idea that art has conveyed scientific concepts to the public... but you’ve heard all that before. This is one of several Bakker images included within the exhibition discussed in this article. Image: (c) Robert Bakker.

Robert Bakker’s 1969 sprinting Deinonychus, produced to accompany John Ostrom’s seminal article on this amazing dinosaur. Artistic depictions like this one cement the idea that art has conveyed scientific concepts to the public... but you’ve heard all that before. This is one of several Bakker images included within the exhibition discussed in this article. Image: (c) Robert Bakker.

During the late 1980s and early 90s, a remarkable thing happened. A team at the Natural History Museum of Los Angeles County, assisted by funding from the Natural History Museum Foundation and fronted by guest curator Sylvia J. Czerkas, created a travelling palaeoart exhibition titled Dinosaurs Past and Present. Opening in Los Angeles in February 1986, and accompanied by a symposium that featured talks on dinosaur science and artwork, the exhibition featured a selection of historical illustrations, paintings and sculptures as well as the very best of contemporary work.

The entrance to what might have been the best palaeoart-themed exhibition of all time, as seen at London’s Natural History Museum in late 1990 and early 1991. A better quality version of this photo will be uploaded to TetZoo in the near future (thanks to Spike Ekins for permission). Image: Spike Ekins.

The entrance to what might have been the best palaeoart-themed exhibition of all time, as seen at London’s Natural History Museum in late 1990 and early 1991. A better quality version of this photo will be uploaded to TetZoo in the near future (thanks to Spike Ekins for permission). Image: Spike Ekins.

Accompanying literature does little to shed light on the backstory of the project and how it all came together – maybe this information is out there and I just haven’t seen it – but the final exhibition essentially functioned as a ‘who’s who’ of late 1980s palaeoart. Credits and acknowledgements show that work was loaned from museums and publishers, but overwhelmingly from private collections, mostly those of the artists themselves. This is interesting for several reasons, one being that the curators and organisers clearly had good relations with the artists, another being that the vast majority of iconic 1980s palaeoart was – as of 1986 at least – owned by its creators and not by the sort of wealthy socialites often associated with niche art. Alas poor palaeoart and its lack of reliable patrons (can someone get that printed on a t-shirt please).

After its Los Angeles opening in 1986, the exhibition toured North America, stops including the Smithsonian, the AMNH, the Tyrrell and the Field. It then crossed the Atlantic for showings in Edinburgh, Cardiff and London, where it finished its run in the January of 1991. Sorry, non-English-speaking nations.

London’s Natural History Museum is an amazing building, decorated inside and out with images of plants and animals. The pterosaur at left is one of several visible on the outside of the museum. Images: Darren Naish.

London’s Natural History Museum is an amazing building, decorated inside and out with images of plants and animals. The pterosaur at left is one of several visible on the outside of the museum. Images: Darren Naish.

I didn’t visit London’s Natural History Museum all that often during my teenage years, but I did get to go there several times as special birthday trips. It was on one of these that – in 1990 – I got to see Dinosaurs Past and Present. This was a total accident, by the way, and not due to clever planning. I didn’t even know about the exhibition beforehand, or that it would be on show during my visit.

Two accompanying volumes – edited by Sylvia J. Czerkas and Everett C. Olson – were produced for the exhibition (Czerkas & Olson 1987a, b). They’re must-haves for serious students of palaeoart, containing many articles that provide invaluable background, discussion and commentary. Those by Mark Hallett, Greg Paul and Dale Russell are especially good. They also, it has to be said, contain several articles that are less valuable, are not especially relevant to palaeoart, and could well have appeared elsewhere.

The covers of volumes I and II of  Dinosaurs Past and Present . Both were initially published (in 1987) in hardback, and later (1989) released as softback. My copies are softbacks. Images: Natural History Museum of Los Angeles County/University of Washington Press.

The covers of volumes I and II of Dinosaurs Past and Present. Both were initially published (in 1987) in hardback, and later (1989) released as softback. My copies are softbacks. Images: Natural History Museum of Los Angeles County/University of Washington Press.

An introduction by John M. Harris (1987) provides some background to the exhibition and its accompanying symposium. Harris’s article, incidentally, includes that specific statement where ‘reconstruction’ and ‘restoration’ are explained to mean different things (p. 4), so it might be this article – rather than the one by Mark Hallett in the same volume – that is best cited as the origin of this concept.

As one of the few people who (a) is of the right age and (b) writes regularly about palaeoart, I’ve been feeling increasingly duty-bound to discuss my recollections of seeing Dinosaurs Past and Present in London in 1990. What, then, do I remember?

First things first: photos were not allowed. You might like to decide among yourselves whether this is a good thing or not. So, despite having a camera with me (a Canon 35 mm of some sort, I think) I didn’t take any photos, except a few sneaky, illegal shots of Stephen Czerkas’s Allosaurus model. More on that model below.

Here’s the only photo I’ve seen that shows what the exhibition looked like. Behind the Czerkas  Allosaurus , framed illustrations on the wall include pieces by Ken Carpenter (at left) and Greg Paul (at right). I can definitely see Paul’s  Styracosaurus  vs  Albertosaurus  image. A better quality version of this photo will be uploaded to TetZoo in the near future (thanks to Spike Ekins for permission). Image: Spike Ekins.

Here’s the only photo I’ve seen that shows what the exhibition looked like. Behind the Czerkas Allosaurus, framed illustrations on the wall include pieces by Ken Carpenter (at left) and Greg Paul (at right). I can definitely see Paul’s Styracosaurus vs Albertosaurus image. A better quality version of this photo will be uploaded to TetZoo in the near future (thanks to Spike Ekins for permission). Image: Spike Ekins.

As goes the exhibition as a whole, I recall it being in one of the galleries that are perpendicular to the marine reptiles corridor (though I could be wrong). The illustrations were hung on white panelling erected to cover the walls (as confirmed by the photos here, by Spike Ekins), there also being a white partitioning wall, reaching to chest- or shoulder-height, erected along the middle of the gallery. Illustrations were hung from either side of this partition as well. There must have been a shop somewhere, since I distinctly recall the two volumes of Dinosaurs Past and Present being on sale. I was able to buy one of them during my visit (I went for Volume II), eventually obtaining Volume I some years later on a separate trip. I recall these books as being cripplingly expensive and my buying of Volume I consuming essentially all of my Natural History Museum trip budget, so it’s embarrassing now to see how (relatively) inexpensive they actually were.

My copies of the  Dinosaurs Past and Present  books still contain their original price labels. Oh, not as expensive as I remembered, then. Whatever. Image: Darren Naish.

My copies of the Dinosaurs Past and Present books still contain their original price labels. Oh, not as expensive as I remembered, then. Whatever. Image: Darren Naish.

Volume I of Dinosaurs Past and Present includes a checklist of all the art featured in the exhibition. No less than 144 pieces were included, the majority being paintings and drawings. They were arranged chronologically, pieces by the likes of Benjamin Waterhouse Hawkins, Charles Knight and Richard Swan Lull being nearest the entrance and the more contemporary work of Mark Hallett, Ely Kish, William Stout, Greg Paul and so on being encountered later on.

Charles Knight’s  Dryptosaurus  painting of 1897 was included in the exhibition and was loaned from the American Museum of Natural History. It’s 58 cm long, 40 cm tall. Image: public domain, wikipedia (original   here  ).

Charles Knight’s Dryptosaurus painting of 1897 was included in the exhibition and was loaned from the American Museum of Natural History. It’s 58 cm long, 40 cm tall. Image: public domain, wikipedia (original here).

Among the older works, I recall seeing the 1879/1880 ‘Pleasures of Science’ piece by Arthur Lakes, and the Benjamin Waterhouse Hawkins depictions of Iguanodon, Hylaeosaurus and Megalosaurus. Weirdly, I have no firm recollection of seeing the works of Charles Knight, Rudolph Zallinger or Peter Zallinger, which is distressing given how influential they’ve been.

Rudolph Zallinger’s incredible mural at Yale’s Peabody Museum is virtually never shown in published photos, and most of the images you’ve seen - like this one - are from the prototype ‘Study’, in which the animals look quite different. Image: (c) Rudolph Zallinger/Peabody Museum of Natural History, Yale University.

Rudolph Zallinger’s incredible mural at Yale’s Peabody Museum is virtually never shown in published photos, and most of the images you’ve seen - like this one - are from the prototype ‘Study’, in which the animals look quite different. Image: (c) Rudolph Zallinger/Peabody Museum of Natural History, Yale University.

In fact, most of what I remember is decidedly contemporary. Of the Mark Hallett originals on show, I remember the montage images that show representative members of assorted clades. My strongest memory is of the ornithopod group, partly because I recall noticing that a hadrosaur originally meant to be a Tsintaosaurus (and shown with the classic, erroneous ‘unicorn’ crest*) had been ‘corrected’ such that it was now an Edmontosaurus. Hallett’s ‘Iguanodon studies’ is also allotted to memory (perhaps because I always liked his idea that iguanodontian beak tissue might have a serrated, pseudotoothed look), as well as his large Morrison Formation panorama of 1975.

* Tsintaosaurus, it turns out, was neither unicorn-crested, nor flat-headed (as has been argued), but instead equipped with a tall, bulbous crest (Prieto-Márquez & Wagner 2013). No more phallic jokes then.

Mark Hallett’s ‘Crossing the Flats’ was at the exhibition. It’s a big piece, more than 1.2 m long. Today we don’t think that  Mamenchisaurus  looked quite like this (Mark was basing his reconstruction on the idea that  Mamenchisaurus  was a diplodocid). This painting always interested me because of the bipedal, narrow-chord pterosaurs as much as the sauropods. Image: (c) Mark Hallett.

Mark Hallett’s ‘Crossing the Flats’ was at the exhibition. It’s a big piece, more than 1.2 m long. Today we don’t think that Mamenchisaurus looked quite like this (Mark was basing his reconstruction on the idea that Mamenchisaurus was a diplodocid). This painting always interested me because of the bipedal, narrow-chord pterosaurs as much as the sauropods. Image: (c) Mark Hallett.

I also recall Ken Carpenter’s illustrations of Sauropelta and ‘Velociraptor’ (proof that Ken was following Greg Paul’s nomenclature; the ‘Velociraptor’ here actually being Deinonychus).

The Greg Paul Years. I make no secret of the fact – it’s mentioned in most books and articles I’ve written or contributed to on dinosaurs and palaeoart (Naish 2009, Conway et al. 2012, Naish & Barrett 2018) – that I consider Greg Paul one of the most influential and important of palaeoartists, and this is true even if you disagree with various of his contentions. Indeed, Greg’s significance in palaeoart is demonstrated by the fact that a full 24 of his pieces were included within the exhibition, more than any other artist.

Greg Paul’s 1987 article from  Dinosaurs Past and Present, Volume II  remains one of the best guides to the life appearance of extinct archosaurs, even though it’s now substantially dated. John Conway and I aim to produce a volume that ‘replaces’ it at some point; meanwhile there’s Mark Witton’s excellent  The Palaeoartist’s Handbook  (which will be reviewed here at TetZoo soon). Image: Darren Naish.

Greg Paul’s 1987 article from Dinosaurs Past and Present, Volume II remains one of the best guides to the life appearance of extinct archosaurs, even though it’s now substantially dated. John Conway and I aim to produce a volume that ‘replaces’ it at some point; meanwhile there’s Mark Witton’s excellent The Palaeoartist’s Handbook (which will be reviewed here at TetZoo soon). Image: Darren Naish.

Unsurprisingly, then, my main recollections concern his pieces. His black and white artwork is mostly small (say, 40 x 40 cm or so), numerous corrections and edits being visible where they were made either with paper whitener (we tend to call it tipex in the UK due to one specific brand) or with pieces of card that have been stuck over the relevant areas. I distinctly recall the tail of one of the tyrannosaurs in his ‘Monoclonius albertensis Fending Off Albertosaurus libratus’ (“Monoclonius albertensis” = Styracosaurus) revealing obvious signs of having its angle corrected (a familiar issue to those who’ve drawn dinosaurs). His ‘What Happens When Apatosaurus ajax Seeks Aquatic Refuge from Allosaurus fragilis’ originally had the riverbed too high, a correctional piece of card allowing him to position it further down the canvas, thereby allowing more space for the swimming sauropod’s feet.

I’m pretty sure I also recall looking at his iconic Giraffitan scene, at his G. brancai muscle study, and at the scene showing Iguanodon and Mantellisaurus (I. atherfieldensis at the time) foraging alongside one another.

Greg Paul’s art is among the most influential dinosaur-themed palaeoart ever produced. Today we know that his feathered non-bird theropods aren’t feathery enough but, hey, you have to start somewhere. This painting was featured in the exhibition. Image (c) Greg Paul.

Greg Paul’s art is among the most influential dinosaur-themed palaeoart ever produced. Today we know that his feathered non-bird theropods aren’t feathery enough but, hey, you have to start somewhere. This painting was featured in the exhibition. Image (c) Greg Paul.

I also recall a few colour Greg Paul paintings. I definitely remember the ‘Resting Velociraptor antirrhopus Pair’ painting, perhaps in part because Greg’s ideas on feathered dinosaurs, and his taxonomic argument that Deinonychus should be considered a species of Velociraptor (which he’s since abandoned), were novel to me at the time. I think that the Allosaurus vs Diplodocus scene was there but my recollection is hazy. I’m far more confident about seeing Paul’s painting of the Pentaceratops herd. I remember it being pretty big, a metre or so in width. To, again, my disappointment, I have no recollection whatsoever of seeing the ‘Tyrannosaurus torosus in a Fast Run’ (for “Tyrannosaurus torosus” read Daspletosaurus). Shocking, because this is another iconic image of the Dinosaur Renaissance.

Greg Paul’s running  Daspletosaurus  - here on the cover of   Lauber (1989)   - is one of his most famous paintings, yet I don’t remember seeing it. Oh well. Image: Darren Naish.

Greg Paul’s running Daspletosaurus - here on the cover of Lauber (1989) - is one of his most famous paintings, yet I don’t remember seeing it. Oh well. Image: Darren Naish.

Enough with the Greg Paul. What else do I remember? I do distinctly remember seeing colour pieces by John Gurche. Gurche has the most incredible style, his colour pieces looking like photos and very often including a marked contrast between a brightly lit, extremely sunny portion and a pitch-black area of deep shadow.

Some of Gurche’s paintings are surprisingly small. On seeing the 1982 Archaeopteryx piece, I was struck by its small size (around 20 x 30 cm). I don’t remember seeing his 1985 Daspletosaurus vs Styracosaurus piece, but it was definitely there. Again, what is wrong with my memory?

A John Gurche painting of 1985 appeared on the cover of an especially famous and influential book (  Bakker 1986  ). Image: Penguin Books.

A John Gurche painting of 1985 appeared on the cover of an especially famous and influential book (Bakker 1986). Image: Penguin Books.

On that note: bizarrely, I don’t remember seeing pieces by Doug Henderson, Robert Bakker or Ely Kish, even though they were definitely there too. Weird. Swiss cheese memory. Nor do I remember seeing the original egg tempera study of Zallinger’s Yale mural, Hallett’s ‘Crossing the Flats’, or a hundred other significant pieces that were there. The more I think about this, the angrier I become. Maybe all that drinking and recreational drug use is to blame. Ha ha, kidding, kidding, kidding.

For its stint at the Royal Museum of Scotland in Edinburgh, publicity for Dinosaurs Past and Present included this photo-shoot, involving the late palaeontologist Beverly Halstead and Ron Seguin’s troodontid and dinosauroid models. Halstead is at far right. I don’t know if Halstead is goofing around (he has his arm around the dinosauroid’s shoulders), but it looks like he is. Image: (c)  New Scientist .

For its stint at the Royal Museum of Scotland in Edinburgh, publicity for Dinosaurs Past and Present included this photo-shoot, involving the late palaeontologist Beverly Halstead and Ron Seguin’s troodontid and dinosauroid models. Halstead is at far right. I don’t know if Halstead is goofing around (he has his arm around the dinosauroid’s shoulders), but it looks like he is. Image: (c) New Scientist.

Finally, what about models? Yes, the dinosauroid and the accompanying Stenonychosaurus were both there, and oh does it hurt to not have any photos of them. You’ve surely seen colour photos of both of these models (created by Ron Seguin in co-operation with Dr Dale Russell): they’re reproduced in many books. Large potted plants (including parlour palms and cycads) were arranged around the models to create a slightly greener vibe. You can see all of this in the black and white photo reproduced above, taken to show the late Beverly Halstead with the models while Dinosaurs Past and Present was on show in Edinburgh.

One of several books that were highly inspirational if you encountered them at the right time.   Wallace (1989)   includes both exciting artwork as well as cutting-edge news ( Protoavis ?! Wtf!, I thought). And on the cover? Yes, it’s Czerkas’s allosaur, portrayed as a dark and sinister predator: “ the Darth Vader of animals ”, to quote John Conway. Image: Darren Naish.

One of several books that were highly inspirational if you encountered them at the right time. Wallace (1989) includes both exciting artwork as well as cutting-edge news (Protoavis?! Wtf!, I thought). And on the cover? Yes, it’s Czerkas’s allosaur, portrayed as a dark and sinister predator: “the Darth Vader of animals”, to quote John Conway. Image: Darren Naish.

Nearby, Stephen Czerkas’s half-life-size Allosaurus model was on show. As mentioned above, this is the one exhibit I have photos of (though I can only find one of my photos today, dammit). At half life-size, it stands perhaps 1.5 m at the top of the head. It was browner than I always imagined: based on the cover of Joseph Wallace’s 1989 The Rise and Fall of the Dinosaur* – an influential book for the young Darren Naish – I always thought it was dark grey or black. Like all of Czerkas’s models, it included a spectacular and really impressive amount of detail and was a joy to look at.

* I find it a bit annoying that Steve Brusatte’s new book has essentially the same title. Steve didn’t know of Wallace’s book until I told him about it, so not his fault. Did no-one think to say anything?

Stephen Czerkas’s brilliant  Allosaurus  model at the Natural History Museum, London (as demonstrated by the accompanying stonework) in 1990. Note that the model is in a different position relative to where it is in Spike’s photos shown above. Image: Darren Naish.

Stephen Czerkas’s brilliant Allosaurus model at the Natural History Museum, London (as demonstrated by the accompanying stonework) in 1990. Note that the model is in a different position relative to where it is in Spike’s photos shown above. Image: Darren Naish.

Several smaller models, made by Stephen and his wife Sylvia Czerkas, were also on show and on top of white, rectangular display stands. I remember Sylvia’s hatching Protoceratops and, I think, Stephen’s 1986 Stegosaurus, controversially constructed with the single row of plates that Stephen thought correct (Czerkas 1987).

For completist reasons I should note that I did get to meet Stephen, once, at the Denver 1999 Society of Vertebrate Paleontology meeting. We spoke briefly about ‘Archaeoraptor’, later unveiled as a composite but thought by Czerkas at the time to be a long-tailed early bird, but I never did get to talk to him about his art.

Anyway, that about wraps things up. Clearly, my memory of a truly momentous and significant exhibition is alarmingly deficient, and the fact that I don’t really have any photographic record of what I saw just makes it worse. What I do remember was, however, thrilling. There have, of course, been a good number of palaeoart-themed exhibitions since 1991, but I think it’s fair to say that none have been as grand, culturally significant or momentous as Dinosaurs Past and Present. Could its like ever occur again? Maybe time will tell, or maybe I’m nostalgic for a Golden Age which has long since passed.

What has happened in the world of palaeoart since the days of Dinosaurs Past and Present? Quite a lot… Image: Darren Naish.

What has happened in the world of palaeoart since the days of Dinosaurs Past and Present? Quite a lot… Image: Darren Naish.

We will finish with the usual reminder that I basically have an infinite number of articles I want to write for TetZoo, but workload and the quest for solvency prevents me from being more productive. If you like what I do, you can help by supporting me at patreon. Thank you!

Palaeoart and changing ideas on the life appearance of Mesozoic dinosaurs have been covered at TetZoo many times over the years. Many of the older articles are now useless because malfunction at the hosting sites has removed their illustrations. Anyway, see…

Refs - -

Bakker, R. T. 1986. The Dinosaur Heresies. Penguin Books, London.

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

Czerkas, S. A. 1987. A reevaluation of the plate arrangement on Stegosaurus stenops. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present, Volume II. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 82-99.

Czerkas, S. J. & Olson, E. C. 1987a. Dinosaurs Past and Present, Volume I. Natural History Museum of Los Angeles County/University of Washington Press, Seattle and Washington.

Czerkas, S. J. & Olson, E. C. 1987b. Dinosaurs Past and Present, Volume II. Natural History Museum of Los Angeles County/University of Washington Press, Seattle and Washington.

Harris, J. M. 1987. Introduction. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present, Volume I. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 1-6.

Lauber, P. 1989. The News About Dinosaurs. Bradbury Press, New York.

Naish, D. 2009. The Great Dinosaur Discoveries. University of California Press, Berkeley and Los Angeles.

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

Prieto-Márquez, A. & Wagner, J. R. 2013. The 'unicorn' dinosaur that wasn't: a new reconstruction of the crest of Tsintaosaurus and the early evolution of the lambeosaurine crest and rostrum. PLoS ONE 8 (11): e82268.

Wallace, J. 1989. The Rise and Fall of the Dinosaur. David & Charles, Newton Abbot, London.

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

You don’t have to be an expert on the fossil birds of the Mesozoic Era to know that our knowledge of these animals has increased exponentially in recent years. An extraordinary number of new species have been described, we’ve learnt a great deal about their anatomy thanks to spectacular new fossils – some of which are even preserved in amber – and we’ve gained insights into their behaviour and ecology thanks to discoveries made about their teeth, feathers, stomach contents, phylogeny and the environments in which they lived.

Enantiornithines — here are just a few of them. In-prep montage from my   in-prep textbook  . Image: Darren Naish.

Enantiornithines — here are just a few of them. In-prep montage from my in-prep textbook. Image: Darren Naish.

Among the most important and species-rich of Mesozoic bird groups are the enantiornithines, or ‘opposite birds’, so named because a few aspects of their skeletal anatomy (the way their shoulder and chest bones fit together in particular) are unusual relative to those of modern birds. Enantiornithines are known from rocks worldwide and were present from the start of the Cretaceous until its close 66 million years ago. They might even have evolved in the Jurassic, in which case older members of the group await discovery.

Among the many interesting things discovered recently about enantiornithines is that at least some of them were colonial nesters. In a Naturwissenschaften paper published in 2012, Gareth Dyke, Mátyás Vremir, Gary Kaiser and myself reported a remarkable fossil assemblage: a big, lens-shaped mass of calcareous mudstone (about 80 cm long, 50 cm wide and 20 cm deep) packed with literally thousands of enantiornithine eggshell fragments. A few enantiornithine bones were present as well, but eggshell fragments form 70-80% of the entire mass (Dyke et al. 2012).

Different chunks of the (originally lens-shaped) Oarda de Jos eggshell assemblage. You can see abundant eggshell fragments (a, b) as well as crushed but complete eggs (the image at bottom). Scale bars = 1 cm. Image: Vremir, Dyke  et al .

Different chunks of the (originally lens-shaped) Oarda de Jos eggshell assemblage. You can see abundant eggshell fragments (a, b) as well as crushed but complete eggs (the image at bottom). Scale bars = 1 cm. Image: Vremir, Dyke et al.

This eggshell mass was found at Oarda de Jos near Sebeş, Transylvania, western Romania and comes from the latest Cretaceous Sebeş Formation (Dyke et al. 2012). Pleurodire turtles, azhdarchid pterosaurs and such dinosaurs as ornithopods and the peculiar Balaur bondoc (which is probably a flightless member of the bird lineage, not a dromaeosaur as originally proposed: see Cau et al. 2015) also come from the Sebeş Formation. I wrote about the discovery and initial interpretation of the eggshell assemblage at TetZoo ver 3, though (sigh) it now appears without any of its images.

The Romanian maniraptoran theropod  Balaur bondoc , originally published as a dromaeosaurid, has a number of features which indicate that an avialan position might be more likely, and this is the position it has occupied in several phylogenetic analyses. If it really is an avialan, it has to be interpreted as a big, secondarily flightless member of the group. We explored this idea in   Cau  et al . (2015)  . Image: Emily Willoughby.

The Romanian maniraptoran theropod Balaur bondoc, originally published as a dromaeosaurid, has a number of features which indicate that an avialan position might be more likely, and this is the position it has occupied in several phylogenetic analyses. If it really is an avialan, it has to be interpreted as a big, secondarily flightless member of the group. We explored this idea in Cau et al. (2015). Image: Emily Willoughby.

As described in our 2012 paper, the assemblage is a fairly big deal, since it means that colonial nesting was practised by at least some enantiornithine species, is not unique to crown-birds, and evolved in birds more stem-ward than other colonially nesting birds (Naish 2014).

But as revealed in our new paper, published in Scientific Communications this past week and led by Mariela Soledad Fernández, things turn out to be a bit more complex than we originally thought. Rather than consisting of enantiornithine eggshell fragments and bones alone, the assemblage actually contains eggshell fragments (and probably bones) of several different, additional animal groups. Namely, gekkotan lizards, crocodylomorphs, and a bird different from the enantiornithine otherwise so well represented in the assemblage (Fernández et al. 2019).

Examined under microscopes (A-B show thin-sections viewed with a standard light microscope; C-D views from the SEM), eggshell fragments look like this. These images show the crocodylomorph eggshell in the assemblage. Image:   Fernández  et al . (2019)  .

Examined under microscopes (A-B show thin-sections viewed with a standard light microscope; C-D views from the SEM), eggshell fragments look like this. These images show the crocodylomorph eggshell in the assemblage. Image: Fernández et al. (2019).

It should be noted to start with that enantiornithine remains dominate the assemblage by far, around 70% of the sampled eggshell fragments belonging to that group (and presumably to the same one species). Sadly, the gekkotan and crocodylomorph eggshell fragments aren’t informative enough to tell us anything particularly interesting about the species concerned, other than that their remains are present. The Oarda de Jos crocodylomorph eggshell is different in thickness and microscopic surface texture from crocodylomorph eggshell fragments reported from the Upper Cretaceous of the USA and Brazil and is most similar to fossil eggs from the Eocene of Colorado, called Krokolites wilsoni (Hirsch 1985). But we don’t have a good handle on what sort of crocodylomorph we’re talking about. After the enantiornithine, its eggs are the most abundant in the sample, forming about 28% of the assemblage. Those artistic reconstructions previously created for the location are thus in error: they really should have at least a few crocodylomorphs in view.

Remember this scene? Produced in 2012 by   Julio Lacerda  , it depicts the possible appearance of the enantiornithine nesting colony we infer for the locality. Maybe some of the colony did look like this. But it now seems that a few crocodylomorphs and the odd lizard were in the immediate area as well. Image:   Julio Lacerda  . UPDATE: this is a horribly lo-res version of the image, I aim to publish a better one in time.

Remember this scene? Produced in 2012 by Julio Lacerda, it depicts the possible appearance of the enantiornithine nesting colony we infer for the locality. Maybe some of the colony did look like this. But it now seems that a few crocodylomorphs and the odd lizard were in the immediate area as well. Image: Julio Lacerda. UPDATE: this is a horribly lo-res version of the image, I aim to publish a better one in time.

The gekkotan eggshell pieces – they have an eggshell morphology termed ‘geckoid’ – have features in common with the eggs of modern geckos, and hence were presumably produced by crown-geckos (Fernández et al. 2019). Beyond that, we can say no more. Less than 1% of the eggshell in the assemblage comes from this lizard, so we’re not saying that its eggs are abundant in the sample.

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

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

Similarly, the second bird in the assemblage appears (from microscopic details of eggshell anatomy) similar to crown-birds – more so than to enantiornithines – but cannot be matched with any specific bird group and might represent something new (Fernández et al. 2019). So, we don’t know exactly what sort of bird we’re dealing with. Again, less than 1% of the eggshell in the assemblage is of this type, so it’s rare in our sample.

Piecing all of this together, what does it all mean? If our enantiornithine eggshell assemblage doesn’t involve enantiornithines alone, but also includes the remains of a second bird, a lizard and a crocodylomorph… do we have an example here of a multi-species nesting colony, perhaps one involving so-called parasitic nesting or even communal or cooperative nesting?

Several examples of this sort of thing are known for the modern world. They involve turtles laying their eggs at the edges of crocodylian nests, South American geckos which lay their eggs within the nests of cormorants and gulls, and seemingly harmonious nesting associations involving rheas, tinamous and penguins. Maybe the behaviours listed here occurred in Maastrichtian Romania. Perhaps the gekkotan lizard concerned was cheekily laying its (hard-shelled, perhaps sticky-shelled) eggs at the edges of enantiornithine nests, and maybe the mystery bird and crocodylomorph were non-threatening enough to be tolerated, their nests perhaps being close to those of the abundant enantiornithines (Fernández et al. 2019).

Opportunistic, parasitic, co-operative and harmonious nesting associations exist in the modern world; here’s an example where raptor nests (in this case, that of an Osprey  Pandion haliaetus  pair) invite the association of passerines, herons and others. Cases like this could well have existed in the Cretaceous. Image: D’Ami  et al . (1969).

Opportunistic, parasitic, co-operative and harmonious nesting associations exist in the modern world; here’s an example where raptor nests (in this case, that of an Osprey Pandion haliaetus pair) invite the association of passerines, herons and others. Cases like this could well have existed in the Cretaceous. Image: D’Ami et al. (1969).

It may also be that the association we report is not quite as interesting as just described, but more to do with geological and hydrodynamic processes. That is, that the eggshell fragments and bones concerned became associated due to their presence in the same general floodplain area, their remains becoming mixed together by the actions of floodwater and not being all that informative as goes behaviour and ecology. Even if this is true, however, we can at least say that these animals were nesting in the same area and environment, and seemingly in proximity. That alone is interesting, and there are indications from elsewhere in the Cretaceous fossil record that it might have been a fairly regular occurrence.

Our paper is open access (OA) and available here; it’s one of several technical papers I hope to see published in the year. On that note, here’s a reminder that I’m not a salaried academic researcher and that any contribution I make to the technical literature is done in my own time. Thanks to those who support me via pledges at patreon, and please consider doing so if you don’t already.

For previous TetZoo articles relevant to Mesozoic birds, the Late Cretaceous animals of Romania and other relevant issues, see…

While compiling this list I’ve discovered that essentially ALL of the TetZoo articles on enantiornithines are hosted at sites (ScienceBlogs and SciAm) that have stripped them of their original images. I must therefore make some effort to republish them here, with their pictures. Great, more stuff to do.

Refs - -

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

D’Ami, R. D., Invernici, F. & Quochi, G. 1969. Animals of Lake and Marsh. Casa Editrice AMZ, Milan.

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

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.

Hirsch, K. F. 1985. Fossil crocodilian eggs from the Eocene of Colorado. Journal of Paleontology 59, 531-542.

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

TetZoo Bookshelf, February 2019, Part 1

There are a lot of books that require review at TetZoo. Let’s see how many I can get through right now…

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Michael J. Everhart’s Oceans of Kansas: A Natural History of the Western Interior Sea, Second Edition

Those of us interested in Mesozoic animals frequently lament the lack of any good, especially useful books that review our knowledge of Mesozoic marine life. But it’s not all bad, since we at least have this wonderful, weighty, very detailed volume. Oceans of Kansas is by far the best there is, and it’s a must-have. Ok, the limitation is that it’s devoted, of course, to Late Cretaceous North American seas alone – specifically, to the Western Interior Sea that once extended north-south across the whole of North America – but this is no fault of the book but its main feature. Indeed, Oceans of Kansas is successful enough that this is its second edition, and it’s newly expanded, revised and augmented, so much so that those owning the first, 2005 edition should consider buying this one too.

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Everhart devotes chapters to sharks, bony fishes, turtles, the various plesiosaur groups, mosasaurs, pterosaurs and dinosaurs as well as invertebrates, the history of our discovery of the Western Interior Sea and more. At 427 pages and about 3.5 cm in thickness, this book contains a huge amount of information. It feels like the compilation of a dedicated person’s life’s work, and I hope the author doesn’t mind me saying that.

Excellent colour photos and numerous diagrams and other illustrations appear throughout. Many of these images are rare or novel and there’s much here not available elsewhere. A colour plate section features art by the late, great Dan Varner, a friend and correspondent I admired and liked a great deal. Dan’s paintings are looking a bit dated now in view of recently acquired information on mosasaur anatomy and we’d all love to know what Dan would have done with them had he the chance (he died in 2012). Regardless, they’re fantastic pieces of work.

One of Dan Varner’s greatest illustrations (in my opinion): the shark  Cretoxyrhina  takes out a  Tylosaurus . This and many other Varner pieces feature in  Oceans of Kansas . Image: (c)   Dan Varner/Oceans of Kansas  .

One of Dan Varner’s greatest illustrations (in my opinion): the shark Cretoxyrhina takes out a Tylosaurus. This and many other Varner pieces feature in Oceans of Kansas. Image: (c) Dan Varner/Oceans of Kansas.

Everhart’s text combines historical review with discussions of what we know, or think we know. There are also numerous personal anecdotes about how various specimens were discovered, recovered and interpreted as well as discussions on how relevant ideas on Cretaceous marine life have developed or changed during the time in which the author has been involved. As he explains, some ideas (those on mosasaur skin and physiology, for example) have undergone substantial revision since he published the book’s first edition in 2005. It’s this personal touch that I enjoy the most.

I say again that Oceans of Kansas is absolutely a must-have if you’re interested in Mesozoic marine life. In my work as a vertebrate palaeontologist and researcher I’ve consulted it hundreds of times, and – like all very good books – it’s not just a great source of information but also a joy to look at.

Michael J. Everhart. 2017. Oceans of Kansas: A Natural History of the Western Interior Sea, Second Edition. Indiana University Press (Bloomington, Indiana), pp. 427. ISBN 978-0-253-02632-3. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Katrina van Grouw’s Unnatural Selection

Much of the fandom that developed around Katrina van Grouw’s 2013 The Unfeathered Bird did so because of its spectacular, beautiful illustrations (van Grouw 2013). Van Grouw’s newest book – 2018’s Unnatural Selection – has a similar format and design and is also spectacularly well illustrated; it is, however, an important work of literature, not of illustration or art alone. I’ve already said quite a fair bit about this book – I wrote a long article on its contents, published before it was out – back in May 2018 at ver 2, so will be brief here.

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Divided into four main sections (Origin, Inheritance, Variation and Selection), Unnatural Selection describes and celebrates selective breeding and its results. We’ve bred animals with all manner of anatomical novelties not present in their wild ancestors, and – as van Grouw explains right at the start of the book – have ended up with domestic hybrids and variants that are so modified that ancestry has been obscured, or are hybrid mish-mashes of more than one species.

Unnatural Selection  showcases an extraordinary number of unusual domestic animals (these are all domestic pigeon breeds), but the main theme is the evolutionary plasticity, and potential, that human selective breeding has created in these species. Image:   van Grouw (2018)  .

Unnatural Selection showcases an extraordinary number of unusual domestic animals (these are all domestic pigeon breeds), but the main theme is the evolutionary plasticity, and potential, that human selective breeding has created in these species. Image: van Grouw (2018).

There’s a huge quantity of information here on the diversity of domestic animals and the backstories to them. Van Grouw talks about specific breeders and what they aimed to do with particular animal breeds, as well as the (sometimes erroneous) ideas on why certain breeds are the way they are. But… this isn’t really what the book is about. The main thrust here, instead, is to highlight how selective breeding has enhanced and modified the variation and plasticity present ancestrally in the animals concerned, and on how selective breeding is super-rapid, human-controlled evolution, echoing, mirroring or replicating that which has occurred outside of human influence.

Geese (wild and domesticate) from    Unnatural Selection   . This panel combines Katrina’s illustrations with Natee Himmapaan’s amazing skills in writing and labelling. Credit: Katrina van Grouw/  van Grouw (2018)  .

Geese (wild and domesticate) from Unnatural Selection. This panel combines Katrina’s illustrations with Natee Himmapaan’s amazing skills in writing and labelling. Credit: Katrina van Grouw/van Grouw (2018).

If you’re interested in the diversity of domestic animals, in evolution, or in anatomy, you absolutely must get hold of this remarkable book, and all the more so and if you enjoyed The Unfeathered Bird.

Katrina van Grouw. 2018. Unnatural Selection. Princeton University Press (Princeton and Oxford), pp. 284. ISBN 978-0-691-15706-1. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

 Jonathan Losos’s Improbable Destinies: Fate, Chance, and the Future of Evolution

Why certain things have happened in evolutionary history, why certain other things haven’t happened, and what things might be like had events took a different turn has, as you’ll know, been a popular area of discussion in this neck of the woods since whenever. Broadly speaking, I have SpecBio in mind: big-brained dinosaurs, an Earth ruled by giant lizards, that sort of thing. If this subject interests you, you absolutely have to read Jonathan Losos’s Improbable Destinies. Inspired (in part) by Gould’s argument in Wonderful Life that evolutionary history is and was dependent on contingency and not destiny, and with something of a herpetological bias linked to his specialisation as a student of anoles, Losos explores the Park Grass Experiment, guppy and stickleback variation, fruit fly studies, disease biology and more.

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My own biases mean that I especially enjoyed his take on dinosauroids (“… to my surprise … The dinosauroid hypothesis was alive and thriving in cyberspace”; p. 323), both the Dale Russell version and the Kosemen/Roy one (yes, there is an illustration), and I think I recall providing him with some of the relevant literature (Naish 2008). The same chapter also includes a discussion of Perry the Platypus and platypusoids (yes, really), E. O. Wilson’s thoughts on extraterrestrials, and, ugh, Simon Conway Morris’s take on the whole thing.

Marlin Peterson’s re-drawing of one of the Kosmen/Roy dinosauroids, from Jonathan Losos’s  Improbable Destinies . Image:   Losos (2017)  .

Marlin Peterson’s re-drawing of one of the Kosmen/Roy dinosauroids, from Jonathan Losos’s Improbable Destinies. Image: Losos (2017).

Losos’s writing is fun and eloquent and the book is really well designed. References and detailed footnotes are included, and the black and white illustrations by Marlin Peterson are really nice. If you enjoy reading this blog, you will like this book. UPDATE: hilariously, I just discovered that I reviewed this book before, here at TetZoo ver 2 in December 2017. I covered exactly the same points because my brain only works one way.

Jonathan B. Losos. 2017. Improbable Destinies: Fate, Chance, and the Future of Evolution. Riverhead Books (New York), pp. 368. ISBN 978-0-399-18492-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Laufer et al.’s Audubon’s Last Wilderness Journey: The Viviparous Quadrupeds of North America

Most people reading this, especially those interested in the portrayal of animals in art, will be familiar with the several works of naturalist, ornithologist and artist John James Audubon (1785-1851). Audubon is most famous for his grand, lavish The Birds of America, published between 1827 and 1839 and involving years of work, massive investment and hundreds of illustrations. But he should be just as well known for his equally momentous and lavish The Viviparous Quadrupeds of North America.

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This new book – including contributions from nine authors, and edited by Charles T. Butler – reproduces all 150 prints from that work, includes a great deal of material that essentially serves as ‘the making of The Viviparous Quadrupeds of North America’ (actually, this is the exact title of Ron Tyler’s article within the volume), and also features sections of text on the history of conservation and natural history in North America, and on how human-wildlife interactions have changed since Audubon’s time. Footnotes cite the relevant correspondence and literature. These ‘introductory’ sections take us all the way to p. 84, so this is a substantial volume (31 x 26 cm, 280 pp).

As goes the illustrations, the quality of reproduction is outstandingly good. The colours are deep and vibrant, and the level of detail that shows up in magnified sections is impressive. Aubudon was really good at drawing hairs, and my god his skunks, foxes, bears and others clearly involved a lot of work and are quite something to examine in detail. The plates are reproduced in entirety, but magnified sections are also included for some of them as well, and it was these that I enjoyed looking at the most.

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The volume ends with a timeline that explains every step in the creation of The Viviparous Quadrupeds, including reproductions of correspondence and colour maps of Audubon’s journey.

This is a grand, beautiful book that will be appreciated by anyone interested in mammals or natural history more broadly. Those especially interested in Audubon and his work will appreciate the biographical and background information, and those who don’t own previous reproductions of The Viviparous Quadrupeds (I imagine that that’s most of us) will enjoy the excellent reproductions of his art.

Marilyn Laufer, Ron Tyler, Charles T. Butler, Dennis Harper, Daniel Patterson, Sarah Zohdy, Robert A. Gitzen, James B. Armstrong & Christopher A. Lepczyk. 2018. Audubon’s Last Wilderness Journey: The Viviparous Quadrupeds of North America. Jule Collins Smith Museum of Fine Art, Auburn University/D. Giles, London. pp. 280. ISBN 978-1-911282-10-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk (where title is spelt wrong.. I have told amazon), here from the publishers.

John Reilly’s The Ascent of Birds

It has always been a great paradox to me why books on the evolutionary history of birds are so incredibly rare. There is – as you’ll know if you’re at all familiar with books on natural history – a wonderful yet maddeningly frustrating never-ending glut of excellent, glossy, beautiful, horridly expensive books on birds, year on year on year, yet bird evolution just never gets covered. Regular readers will know what I think of Alan Feduccia’s idiosyncratic, biased and utterly misleading writings.

It is a major problem that there has long been a total shortage of reliable books on bird evolution. Alan Feduccia’s books - two of which are shown here - are unreliable and misleading. Images:  Yale University Press (left);   Yale University Press (right) .

It is a major problem that there has long been a total shortage of reliable books on bird evolution. Alan Feduccia’s books - two of which are shown here - are unreliable and misleading. Images: Yale University Press (left); Yale University Press (right).

John Reilly’s volume substantially alleviates that gap in the market (yes, I do know about Gerald Mayr’s books), and it’s really good. Written very much in the format of Dawkins’s The Ancestor’s Tale (which I confess to not liking all that much, ask me if you want to know why), The Ascent of Birds is a very dense, very well informed, mostly up to date tour of the bird family tree, its structure following molecular phylogenies like those of Hackett et al., Jarvis et al., Prum et al. and so on: specifically, he follows the topology published by Jetz et al. (2012). I guess he used Jetz et al. (2012) specifically because it was the most up to date at the time of writing (various of its conclusions have been replaced by those more recent studies). My own (now dated) review of bird evolution (Naish 2012) is cited, but I think only as a source of information on passerine genitalia. Ok.

The excellent cover art - featuring paintings by Jon Fjeldså (whose illustrations appear in several of his own papers) - depicts the cladogram published by Jetz  et al . (2012). Image:   Pelagic Publishing  .

The excellent cover art - featuring paintings by Jon Fjeldså (whose illustrations appear in several of his own papers) - depicts the cladogram published by Jetz et al. (2012). Image: Pelagic Publishing.

Chapters discuss lineages in the order of their branching in the Jetz et al. cladogram.. broadly speaking (there are actually a few deviations). Each combines something on the general history of the lineage before focusing on a story specific to one, some or all members of said lineage. The chapter on albatrosses, for example, examines models of albatross biogeography before discussing dynamic soaring behaviour and olfactory biology while that on corvids mostly focuses on cognitive skills and their evolution. We thus get a broad-brush picture on the group’s evolution before zooming in on something more specific but still relevant to the group’s story as a whole. This works really well, and Reilly focuses on things that are – in my opinion – more interesting than Dawkins did.

References are provided throughout, most chapters contain a diagram or two, and a colour plate section includes images of fossils and living birds relevant to the main prose.

There are a lot of passerine lineages: this simplified cladogram features the major lineages only. It is fitting that about half of Reilly’s  The Ascent of Birds  is devoted to this one group. Image: Darren Naish,   from my textbook project  .

There are a lot of passerine lineages: this simplified cladogram features the major lineages only. It is fitting that about half of Reilly’s The Ascent of Birds is devoted to this one group. Image: Darren Naish, from my textbook project.

Long-time readers will know that I once invested considerable time in a book that, similarly, took its readers on a tour of the modern, molecular avian family tree. It was less prose-based than Reilly’s book, and with more focus on images, and was ultimately abandoned due to lack of interest from publishers (part of the story is discussed here, though the article concerned is now lacking all of its accompanying images). Reilly’s book does what my book should have done, but in a far more detail-oriented fashion, meaning that we finally have a good volume presenting the vast amount of modern work done on bird evolution to those interested. This is a notable achievement and has been well executed.

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Screengrabs from the aborted Naish ‘The Bird Family Tree’ book. Yes, I should pick it up and get it published. No, I can’t do that due to a list of other projects that prevent such action. Images: Darren Naish.

Screengrabs from the aborted Naish ‘The Bird Family Tree’ book. Yes, I should pick it up and get it published. No, I can’t do that due to a list of other projects that prevent such action. Images: Darren Naish.

John Reilly. 2018. The Ascent of Birds: How Modern Science is Revealing Their Story. Pelagic Press (Exeter), pp. 340. ISBN 978-1-78427-169-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Philippe Geniez’s Snakes of Europe, North Africa and the Middle East: A Photographic Guide

Originally published in French, and translated into English by Tony Williams, Geniez’s Snakes is a compact, attractive, well designed and pleasingly thick volume, definitely one to get if you’re a snake specialist. An introductory section covers snake natural history, biology, habitat, the effects of snakebite and classification; the rest of the book – which goes through the species on a family by family basis – covers ‘identification’, ‘range’ as well as diet, reproduction and so on for each species (where available) and is especially good on intraspecific variation and proposed subspecies. Geniez has gone to some trouble to use the newest ideas on taxonomy and phylogeny, the various boxed-out sections providing discussion on genera and their contents. So, you’re going to meet Myriopholis, Spalerosophis, Platyceps and others, if you haven’t already.

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There are already a few (mostly very good) fieldguides to the reptiles of the European field guide region – this typically being taken to include northern Africa and at least part of the Middle East too – so what is it that makes this book different as goes which taxa it includes? Well, there are quite a few species from outside the region usually covered, including a few surprises – too many to discuss here.  I will say that they include the Brahminy worm snake Indotyphlops braminus (yeah, not Ramphotyphlops anymore), California kingsnake Lampropeltis californiae (yup: introduced to the Canary Islands and now abundant there, and also known from Spain), and the as-yet-unnamed Israeli whip snake.

Pages from   Geniez (2018)  . It’s a great-looking book. Image:   weboryx.com twitter account.

Pages from Geniez (2018). It’s a great-looking book. Image: weboryx.com twitter account.

This is not a ‘fieldguide’ in the sense of having plates depicting related species in close proximity. Instead, photos (taken by a long list of different photographers) are distributed throughout the text. But they’re outstandingly good and make this a very colourful, attractive book. Strongly recommended for those of herpetological inclination!

Philippe Geniez. 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press (Princeton and Oxford), pp. 379. ISBN 978-0-691-17239-2. Softback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

 For other TetZoo book reviews and articles relevant to the topics covered here, see…

Refs - -

Geniez. P. 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press, Princeton and Oxford.

Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K. & Mooers, A. O. 2012. The global diversity of birds in space and time. Nature 491, 444-448.

Losos, J. B. 2017. Improbable Destinies: Fate, Chance, and the Future of Evolution. Riverhead Books, New York.

Naish, D. 2008. Intelligent dinosaurs. Fortean Times 239, 52-53.

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

van Grouw, K. 2013. The Unfeathered Bird. Princeton University Press, Princeton and Oxford.

Tetrapod Zoology Is A Teenager Now

Yet again a year has passed and yet again it’s time to review events relevant to the TetZooniverse, so here we are at TetZoo’s 13th birthday (January 21st 2006 being the date on which the blog first came into being). As per usual, I’m going to discuss, comment on or mention all those TetZooniverse things that I deem worthy of discussion, comment or mention. And, as usual, I implore you not to read this if the thought of me writing about my own achievements and adventures sounds arrogant or worth a miss. Still here? Ok, we proceed…

Once again I’m going to begin by reminding you of how it all began, back in 2006… you can see that very first article (should you want to)   here  . Images: Darren Naish.

Once again I’m going to begin by reminding you of how it all began, back in 2006… you can see that very first article (should you want to) here. Images: Darren Naish.

Our story begins in late January 2018. Birds (owls and rollers, among others) were still keeping me busy as goes The Big Book, and academic work on ichthyosaurs, pterosaurs and sea monsters (of the cryptozoological sort) was underway. It’s a reminder (as if another were needed) of how slowly it takes for academic projects to come together to note that none of those projects have yet seen print, more than a year later. Fossil bats – covered only because I suddenly suffered bat guilt – got a little coverage at TetZoo during February. I visited Hill Head, West Wittering and Swanwick to see grebes, divers, waders and gulls. The conditions were mostly cold and windy but clear and sunny, which is great for looking at water- and seabirds.

There are so many great places to see water- and seabirds around the coasts of the UK. One of my favourite haunts at the moment is Swanwick in Hampshire. Teals at left; Brent geese at right. Image: Darren Naish.

There are so many great places to see water- and seabirds around the coasts of the UK. One of my favourite haunts at the moment is Swanwick in Hampshire. Teals at left; Brent geese at right. Image: Darren Naish.

Dinosaurs in the Wild (DITW) opened for its final, London-based run at the start of February. If you’re a regular reader you’ll already be familiar with this interactive, science-based visitor attraction so I won’t be saying much more about it today. If you don’t know what I’m talking about and are curious, see Dinosaurs in the Wild: An Inside View at TetZoo ver 3 and The Last Day of Dinosaurs in the Wild here at ver 4. I did various promotional events for DITW from February to June and met science journalist and author John Pickrell at the exhibition during February. Also in February, I wasted time submitting a grant application which was ultimately unsuccessful – more on that later.

Scenes from Dinosaurs in the Wild. At left, your humble author with John Pickrell (and friend: John is the one not holding the plush tyrannosaur). At right: myself, Siouxsie the  Dakotaraptor , and Chris Packham. Images: Darren Naish, Megan McCubbin.

Scenes from Dinosaurs in the Wild. At left, your humble author with John Pickrell (and friend: John is the one not holding the plush tyrannosaur). At right: myself, Siouxsie the Dakotaraptor, and Chris Packham. Images: Darren Naish, Megan McCubbin.

A TV thing filmed at Colchester Zoo during October 2017 appeared on BBC4, titled ‘The Incredible Science of Temperature’ and presented by Helen Czerski. I was filmed discussing, with Helen, what influence global temperatures have had on the distribution of ectothermic reptiles (qualifier because not all non-bird reptiles are ectothermic), and on the physiology and behaviour of Komodo dragons. It felt, at the time, like a total disaster but I’m pleased to say that the parts of my performance included in the show weren’t too bad (the relevant episode is online here). Fighting blue tits, Leguatia, and an article about Mark Witton’s palaeoart appeared at TetZoo during this part of the year. Remember that TetZoo was still hosted at SciAm at this point.

In March I assisted with a local beach clean event. I don’t talk about these to brag about my public service (heavens, no), but to help maintain awareness of plastic pollution and the problems facing marine environments today, and also to remind those of you reading this that you can help by getting involved with local environmental groups.

Beach cleaning is always such a joy (am being ironic). Here’s a haul after just a few hour’s work. Image: Darren Naish.

Beach cleaning is always such a joy (am being ironic). Here’s a haul after just a few hour’s work. Image: Darren Naish.

March was also the month in which Paul Barrett and I compiled our suggested list of corrections and updates for the second edition of Dinosaurs: How They Lived and Evolved. Again, I don’t need to say much about this here since the story behind the second edition was recently covered here in, err, The Second Edition of Dinosaurs: How They Lived and Evolved. Thanks again to those who provided help in compiling corrections, to those who’ve said positive things about the book and to those who’ve purchased it and helped spread the word about it.

Milo the Neotropical giant toad - at right - now lives at Tet Zoo Towers, but his original home was the BBC Natural History Unit, at BBC Bristol. Like Milo, I spent time at BBC Bristol during 2018. Why? Well… we shall see. Images: Darren Naish.

Milo the Neotropical giant toad - at right - now lives at Tet Zoo Towers, but his original home was the BBC Natural History Unit, at BBC Bristol. Like Milo, I spent time at BBC Bristol during 2018. Why? Well… we shall see. Images: Darren Naish.

I engaged in an online interview with the brilliant Aron Ra who I’ve long been familiar with but had not previously spoken to. We discussed TetZooCon, the Tetrapod Zoology project, my books, serial mis-educator David Peters, and more. The interview – part of Aron’s RaMen series – is online here. I also used our virtual meeting as an excuse to invite Aron as a speaker to TetZooCon 2018 since, by virtue of good timing, he was going to be in the UK at exactly the right time. More on TetZooCon later. Oh: I joined instagram in March, wholly because I read an article by a scientist bemoaning scientists who join instagram. My review of Jonathan Kane et al.’s most worthy book God’s Word or Human Reason? appeared at TetZoo in late March, as did a short review of everything columbiform. By now, the substantial passerine section was being done for The Big Book which surely meant that completion of the enormous bird section was in sight, right?

Work on The Big Book (my in-prep textbook  The Vertebrate Fossil Record ) mostly involved birds across 2018. In cases, as here, I had to revamp some old illustrations. This image shows old (at right) and new images of  Aepyornis … and surely I won’t need to revamp  Aepyornis  AGAIN, right? Some of you will know what I’m getting at. Image: Darren Naish.

Work on The Big Book (my in-prep textbook The Vertebrate Fossil Record) mostly involved birds across 2018. In cases, as here, I had to revamp some old illustrations. This image shows old (at right) and new images of Aepyornis… and surely I won’t need to revamp Aepyornis AGAIN, right? Some of you will know what I’m getting at. Image: Darren Naish.

Sunday Brunch, publishers, duckbills, sexual selection. And so to April. As – once again – part of the whole promotional Dinosaurs in the Wild thing, I appeared on the TV show Sunday Brunch… which I’ve never watched but is, by all accounts, pretty popular. It went well and I got to talk about our current knowledge of Mesozoic dinosaurs a fair bit as well as promote Dinosaurs in the Wild, but what might have been the most memorable section of the interview was at the end where presenter Tim Lovejoy asked what the future might hold in store as goes the evolution of life on Earth. Ever the optimist, I told them about the impending mass extinction and the death of biological diversity at the hands of humanity, and that was that.

A view from the set of  Sunday Brunch , April 2018. Ash, Chrissie and a hatchling  Dakotaraptor  at left. Image: Darren Naish.

A view from the set of Sunday Brunch, April 2018. Ash, Chrissie and a hatchling Dakotaraptor at left. Image: Darren Naish.

Chrissie, Darren and the  Dakotaraptor  hatchling, live on Sunday Brunch on April 8th 2018. Image: (c)  Sunday Brunch /Channel 4.

Chrissie, Darren and the Dakotaraptor hatchling, live on Sunday Brunch on April 8th 2018. Image: (c) Sunday Brunch/Channel 4.

In an effort to get the planned Dinosaurs in the Wild book off the ground, I physically dragged myself from publisher to publisher. So many meetings, and all in vain. While at William Collins, however, I did get to see the complete set of the New Naturalist series and, via publisher Myles Archibald, got to hear the backstory to that Too Big to Walk book… the one by ol’ BJ Ford in which he asserts his uncanny ability to finally get dinosaurs right (they were all aquatic, you see). More on that shortly. I also had a meeting about another planned cryptozoology-based TV series. I can basically guarantee having two or three such meetings about the same sort of project every year. They never go anywhere. Yet.

The Collins New Naturalist books on display at the Collins office in London. Most of us own at least a few of these volumes. Image: Darren Naish.

The Collins New Naturalist books on display at the Collins office in London. Most of us own at least a few of these volumes. Image: Darren Naish.

My article on why duckbilled dinosaurs really shouldn’t be called duckbilled dinosaurs appeared in April. It includes a significant gaff and a follow-up article is due to be published here within the next several weeks. The second part of of my review of the Indiana University Press 2014 hadrosaurs volume was also published in April: the first part of my review is one of those articles that SciAm removed due to perceived problems with image use so isn’t currently online. I’ll republish it here at ver 4, eventually.

Attendees of the Sexual Selection meeting at Chicheley Hall, Buckinghamshire. Image: Rob Knell.

Attendees of the Sexual Selection meeting at Chicheley Hall, Buckinghamshire. Image: Rob Knell.

I attended the Royal Society Sexual Selection meeting at Chicheley Hall in Buckinghamshire in early May and chaired one of the sessions. A TetZoo article about the meeting can be found here at ver 3.

I went to Tintagel, north Cornwall, in May 2018. Here’s a scene of the coastline there, with seabirds. Image: Darren Naish.

I went to Tintagel, north Cornwall, in May 2018. Here’s a scene of the coastline there, with seabirds. Image: Darren Naish.

Naish v Ford. The biggest event of May was the debate with Brian Ford at Conway Hall, hinted at above. It seems that this was actually the official launch for Mr Ford’s book – it’s hard to tell as I didn’t see anybody buy a copy – but it was more like a structured debate, the two of us presenting opposing viewpoints on the palaeobiology of Mesozoic dinosaurs before responding to questions from the floor. Events were reported on twitter via #FordvNaish, and my talk can be seen here online.

At left: the cover slide for the event. At right: protesters outside Conway Hall, as imagined by Gareth Monger. Well done if you recognise the reference. Images: Darren Naish, (c) Gareth Monger.

At left: the cover slide for the event. At right: protesters outside Conway Hall, as imagined by Gareth Monger. Well done if you recognise the reference. Images: Darren Naish, (c) Gareth Monger.

I’ve written about Brian Ford and his ideas at least twice before: both in print for an invited article in Laboratory News (Naish 2012) and at TetZoo ver 3. Needless to say, I think he’s flat-out wrong and also guilty of conspiracy mongering, self-aggrandisement (he has literally compared himself to Galileo) and crankery (“nobody’s smart but me!”), among other things. Evaluating the talks is difficult due to the fact that I’m just about never happy with the presentations I give; in any case, John Conway and I already dissected the event at some length in an episode of the podcast. Having said that, Brian asserted that everyone was getting dinosaurs wrong, and assert is basically all he did. I would think that the mostly negative responses that have been directed toward Mr Ford’s ideas would have been sufficient to make him realise that he could very well be wrong as goes his take on dinosaurs, but… nope… he’s still at it (at least, judging from comments he’s recently been making at ResearchGate). I don’t want to give him more publicity than I already have, but if I need to respond to him again, I will.

Darren Naish (left) and Brian Ford (right). Image: Luke Muscutt.

Darren Naish (left) and Brian Ford (right). Image: Luke Muscutt.

Podcats, Jurassic Park at 25, lego at Metro. Having mentioned Conway, we released a few episodes of the podcast – the Tet Zoo podcats or Tetrapodcats – throughout the first half of the year, but we’re still nowhere near having anything like a regular schedule. Mostly this is because John is incredibly lazy, and unproductive to boot. An added factor is that I ‘accidentally’ cleaved through a buried telephone cable in the garden with a spade one day, but I’m sure that that’s coincidental. The new cover for Dinosaurs How They Lived and Evolved, produced by the excellent Bob Nicholls, was released in May – sooner than expected. More on that below.

In May 2018, I (with my kids Will and Emma) attended the Portsmouth Comic Con, and here are some scenes. Not really relevant to TetZoo, but what the hell. Images: Darren Naish.

In May 2018, I (with my kids Will and Emma) attended the Portsmouth Comic Con, and here are some scenes. Not really relevant to TetZoo, but what the hell. Images: Darren Naish.

Hunting Monsters – the cryptozoology book that I’ve now mentioned here on some number of occasions (Naish 2017) – was mentioned on the TV show Jeopardy in early June, wow. Thanks to whoever it was who brought my attention to this. June 2018 was also the time at which Jurassic Park – a movie you might have seen or heard about – celebrated its 25th birthday. A lot of the fan response to the original Jurassic Park – we see a lot of it in vertebrate palaeontology, in fact it comes around every few years – is, my apologies for seeming rude here, mostly vacuous and doesn’t come from a position all that well informed as goes the backstory and making-of the movie. Make of that what you will; whatever, my three Jurassic Park articles are here, here and here, and my liberal use of Jurassic Park toys reflects my best efforts to circumvent the rules regarding image use at SciAm. For all that, they still objected so much to my use of a photo of an issue of National Geographic that they removed it. So I created my own version of the cover.

Hunting Monsters  appears on  Jeopardy . Image: (c)  Jeopardy , NBC/American Broadcasting Company.

Hunting Monsters appears on Jeopardy. Image: (c) Jeopardy, NBC/American Broadcasting Company.

Another bit of Dinosaurs in the Wild promotion happened in late June when – together with palaeontologist Alessandro Chiarenza and actor Ross Cooper – I got to sit in a studio with Natasha Salmon and build Jurassic World lego, all the while talking about dinosaurs. This was for the Metro newspaper, and what fun it was. I even got to keep some of the lego. I also had an article on dinosaurs – specifically on how our perception of their appearance has changed – appear in the Independent Online in mid July. It’s here.

Playing with  Jurassic World: Fallen Kingdom  lego at the Metro offices. I’m not entirely sure why Natasha is making the face she is. Image: (c) Metro.

Playing with Jurassic World: Fallen Kingdom lego at the Metro offices. I’m not entirely sure why Natasha is making the face she is. Image: (c) Metro.

Eotyrannus 2018. During late June I participated in the 2018 BioTweeps conference (an online, Twitter-based conference in which attendees present their ‘talks’ via a series of threaded tweets), my presentation being ‘Eotyrannus and the History of Tyrant Dinosaurs’ (abstract here). As an invited presenter I got to tweet as much as I wanted to in the allotted 30 minute slot. It was a very rewarding experience (with a massive, global reach, I might add), and my thanks to Dani Rabaiotti and Anthony Caravaggi for getting me involved. You can see my presented tweets here (with an addendum here).

Naish at BioTweeps 2018. Image: Darren Naish.

Naish at BioTweeps 2018. Image: Darren Naish.

Having mentioned Eotyrannus… as some of you know, I am (still) committed to publishing the monograph on this dinosaur, and while I failed to get the damned thing into print in 2018, I promise that this will happen in 2019. It’s not for want of trying: substantial progress was made on the post-review version of the manuscript throughout the year, it’s just that I was never able to commit sufficient time to getting it finished. Silly old me for leaving professional scientific research and becoming a freelancer. My co-author Andrea Cau continues to be a most worthy collaborator.

The  Eotyrannus  holotype, in its storage boxes, at Dinosaur Isle, Sandown, Isle of Wight… as it looked in 2012. One day my work on this dinosaur will be published. Image: Darren Naish.

The Eotyrannus holotype, in its storage boxes, at Dinosaur Isle, Sandown, Isle of Wight… as it looked in 2012. One day my work on this dinosaur will be published. Image: Darren Naish.

I mentioned earlier that I was unsuccessful in obtaining a grant – that’s no big deal, most grant applications in science are unsuccessful. Said grant was an attempt to win the money I require to publish an open-access version of the Eotyrannus monograph. On the advice of colleagues, I tried a different approach and set up a gofundme campaign to raise the money aaaand succeeded in raising it (and a bit extra) in about 5 hours. WOW.

Eotyrannus  at gofundme — SUCCESS. Image: gofundme.

Eotyrannus at gofundme — SUCCESS. Image: gofundme.

Speaking of long-running projects that never seem to reach completion, I and colleagues put what are supposed to be the final touches to a PhyloCode Companion Volume article on theropods during the summer. I’m also part of the team that produced the volume’s sauropod and sauropodomorph articles. But they have a somewhat different fate from the theropod article, that’s all I’ll say for now.

This image has to go somewhere and here is as good a place as any. It’s not made by me, but by naughtyhippo ( if that is her real name ).   Available on merchandise here.   Image: Darren Naish.

This image has to go somewhere and here is as good a place as any. It’s not made by me, but by naughtyhippo (if that is her real name). Available on merchandise here. Image: Darren Naish.

Goodbye SciAm. July was a significant month as goes TetZooniverse things as, on July 25th, I reached a funding goal at patreon. Regular TetZoo readers will know (my god, how many times have l mentioned it?) that I was not happy with the way things were going at SciAm. Their total clamping down on image use and removal of various of my articles was pretty much the last straw: blogging there was proving more trouble that it was worth, despite the major kudos of being attached to the SciAm brand. Achieving the relevant goal meant that I could leave (as a freelancer, I’m dependent on earning from my writing: it is my primary income) and set up shop as an independent blogger once more (harking back to the days of TetZo ver 1, 2006). Despite my unhappiness with SciAm, walking away was not a decision I took lightly. The very last TetZoo ver 3 was published on July 31st, and here we are at ver 4, at the same website at the podcast. Thanks, as ever, to the patreon supporters who made this possible.

Leaving SciAm means that I’m now entirely dependent on   support at Patreon  . What a gamble. As of right now, the site needs updating. Image: Darren Naish.

Leaving SciAm means that I’m now entirely dependent on support at Patreon. What a gamble. As of right now, the site needs updating. Image: Darren Naish.

August began with the taking on of an exciting job in the museums world. It involves an exhibition relevant to our interests, but it won’t go live until 2020 and I won’t be talking about it until then, sorry. I’ll give you a clue and say that it has meant spending time in Cornwall, one of my favourite parts of the UK.  Also during August, I spoke to New Scientist about megalodon in view of a scientific documentary called Meg. I’m not, technically, an expert on megatooth sharks but I have written about them several times: there’s a section in Cryptozoologicon Volume I (Conway et al. 2013) and another – as yet unpublished, of course – in the chondrichthyan section of The Vertebrate Fossil Record.

From time spent in Hay-on-Wye. This is Hay Bluff. Image: Darren Naish.

From time spent in Hay-on-Wye. This is Hay Bluff. Image: Darren Naish.

Articles on the potential domestication potential of non-bird dinosaurs and mastigures (or spiny-tailed agamas) appeared at TetZoo. Together with the family, I visited Hay-on-Wye, the fabled Town of Books of the Welsh borders (and filming location for certain iconic scenes in An American Werewolf in London). There aren’t half as many book shops in Hay-on-Wye as there used to be, but there are enough to make a visit very much worthwhile if you’re a bibliophile. I made a great number of amazing discoveries – beware, there’s a dedicated natural history bookshop (C. Arden) – and am really happy that I finally took the time to go there.

Don’t go to Hay-on-Wye, it will be bad. Image: Darren Naish.

Don’t go to Hay-on-Wye, it will be bad. Image: Darren Naish.

As I mention every now and again, I continue to remain involved in technical scientific research as and when I can, and late August saw the publication of my then-newest paper: a study of pelvic neural anatomy in pterosaurs and what the relevant details might mean for ecomorphology, co-authored with Liz Martin-Silverstone and Dan Sykes (Martin-Silverstone et al. 2018). This was the second 2018 paper that included an analysis of Vectidraco – a small Cretaceous azhdarchoid pterosaur I and colleagues described in 2013 (Naish et al. 2013) – the other being Rachel Frigot’s study of pelvic musculature included within the Geological Society special volume New Perspectives on Pterosaur Palaeobiology (Frigot 2018). A discussion on the new paper was provided here.

At left, an illustration of the  Vectidraco  pelvis, some key features labelled. At right: a pterosaur montage produced for the TetZoo article on pterosaur palaeoneurology. Images: Darren Naish.

At left, an illustration of the Vectidraco pelvis, some key features labelled. At right: a pterosaur montage produced for the TetZoo article on pterosaur palaeoneurology. Images: Darren Naish.

September 2nd was the very final day of Dinosaurs in the Wild. I went on a final tour, attended the staff party, and said all those sad, final goodbyes. All things must come to an end, but hope remains that DITW will be able to open its doors again one day.

Me (at left) with Tim Haines - of Impossible Pictures, Walking With Dinosaurs, Primeval and so on - doing some Dinosaurs in the Wild promotion for radio during May 2018. Image: Darren Naish.

Me (at left) with Tim Haines - of Impossible Pictures, Walking With Dinosaurs, Primeval and so on - doing some Dinosaurs in the Wild promotion for radio during May 2018. Image: Darren Naish.

Dougal Dixon, Crystal Palace, DHLTE: 2nd ed. Early September also brought the devastating news of the calamitous fire at Rio’s National Museum, a blow for human knowledge as a whole but especially for our Brazilian friends and colleagues. While at the museum in 2013, I took some reasonable number of photos, so have been submitting them to Brazilian colleagues. In happier news, September also saw the Dougal Dixon event at London’s Conway Hall (covered here at TetZoo) in which Dougal and I discussed his work and projects in front of a pretty substantial crowd. Original art, models and more were in attendance – it was great.

At Conway Hall, for the  After Man -themed event with Dougal Dixon. Images: Darren Naish.

At Conway Hall, for the After Man-themed event with Dougal Dixon. Images: Darren Naish.

Just a few days later, and I was speaking at another London-based event, this time the Crystal Palace Dinosaurs Days event, part of the Heritage Open Days weekend occurring across the UK. Other speakers were in attendance too, and we had special up-close access to the models. I’m surely repeating myself, as all of this was covered in the December 2018 article Up Close and Personal With the Crystal Palace Dinosaurs. As you may know, part of the reason for the publicity push surrounding the models is that funding is desperately needed for their care and upkeep, and September 2018 saw the launch of a crowd-funded project to get a bridge to the models constructed (said bridge then allowing the access that can otherwise only be obtained on rare occasion). The project reached its target (which was over £70,000) in December, which is excellent and exhilarating news.

Special access to the islands at Crystal Palace, what a treat. Here, I’m talking about what the Victorians thought about ichthyosaurs. For more, see   Up Close and Personal With the Crystal Palace Dinosaurs  . Image: Emma Wain.

Special access to the islands at Crystal Palace, what a treat. Here, I’m talking about what the Victorians thought about ichthyosaurs. For more, see Up Close and Personal With the Crystal Palace Dinosaurs. Image: Emma Wain.

The second edition of Dinosaurs: How They Lived and Evolved – my Natural History Museum book with Paul Barrett (Naish & Barrett 2018) – saw print in September. Again, thoughts on this have already appeared at TetZoo so I’ll avoid repeating things here. A review appeared in The Economist, which is kind of a big deal. Summer 2018 also saw the publication of the Russian edition of the book. It has the text of the second edition but the cover of the first.

Screengrab from the  Economist ’s review of the second edition of   Dinosaurs: How They Lived and Evolved  . The lead quote comes from Bob Nicholls, creator of the fuzzy  Tianyulong  we see at left. Image: (c) Economist.

Screengrab from the Economist’s review of the second edition of Dinosaurs: How They Lived and Evolved. The lead quote comes from Bob Nicholls, creator of the fuzzy Tianyulong we see at left. Image: (c) Economist.

TetZooCon 2018! And so to October, and the TetZoo-themed event of the year, by which I mean TetZooCon. As covered here on the blog, 2018’s TetZooCon was the first two day TetZooCon ever, and by far the biggest and (I think) the best. Speakers and presenters included Ian Redmond, Mark O’Shea, Aron Ra, Dougal Dixon and Gert van Dijk (on stage for a SpecBio discussion), Katrina van Grouw and myself. John Conway led a palaeoart workshop that occurred as a parallel session to some of the talks and I led a post-con fieldtrip to Crystal Palace the day after. A MonsterTalk episode with Blake Smith – done as a piece of TetZooCon promotion but mostly revolving around Jurassic Park – was released in early October (it’s here; episode 172).

Scenes from TetZooCon 2018. At left, Albert Chen discusses TetZoo. At right: a box of Dinosaurs in the Wild plushies. Images: Darren Naish.

Scenes from TetZooCon 2018. At left, Albert Chen discusses TetZoo. At right: a box of Dinosaurs in the Wild plushies. Images: Darren Naish.

Hey, I had to turn down a free, all-expenses-paid birdwatching trip to Cadiz in Spain because it clashed with TetZooCon, talk about bad luck. The success of TetZooCon 2018 means that a 2019 event is just about guaranteed (or, it is now that John and I are back on speaking terms again); it will again be a two-day event but I can’t yet say any more than that. News will be announced as and when I have it.

TetZooCon 2018 included a dedicated SpecBio discussion, and here’s Gert van Dijk talking about his own Furaha project, at said event. Image: Darren Naish.

TetZooCon 2018 included a dedicated SpecBio discussion, and here’s Gert van Dijk talking about his own Furaha project, at said event. Image: Darren Naish.

Dorling Kindersley. It was around this time in the year that I and Chris Barker put the finishing touches to a new kid’s book we’ve put together for Dorling Kindersley, titled Where on Earth? Dinosaurs and Other Prehistoric Life. I wouldn’t mention this (seeing as the book isn’t published yet), but it’s already being advertised.

Screengrabs from the in-press   Where on Earth? Dinosaurs and Other Prehistoric Life  , by Chris Barker and Darren Naish. Images: Barker & Naish/Dorling Kindersley.

Screengrabs from the in-press Where on Earth? Dinosaurs and Other Prehistoric Life, by Chris Barker and Darren Naish. Images: Barker & Naish/Dorling Kindersley.

In fact, I did a huge amount of work for Dorling Kindersley during 2018 as I and colleagues at DK worked hard to overhaul their entire prehistoric animal picture library. One volume that benefited from this overhaul – John Woodward’s The Dinosaurs Book (Woodward 2018) – was published during the year (I was consultant). It should be obvious to anyone paying attention to children’s books on dinosaurs that this overhauling has resulted in a massive improvement as goes the sorts of images the books now contain.

Castle Urquhurt, as photographed from the surface of Loch Ness (a scene from 2014, not 2018). Image: Darren Naish.

Castle Urquhurt, as photographed from the surface of Loch Ness (a scene from 2014, not 2018). Image: Darren Naish.

Loch Ness, 99% Invisible. I’ve mentioned once or twice here my communications with Professor Neil Gemmell and my role in his eDNA-based work on the biology and ecology of Loch Ness. Neil and I met up in 2017 to discuss this work (a 2018 article on the backstory to Neil’s research – one of several – can be found here), and I published two magazine articles on the subject during the year: one in a kid’s magazine in October (Naish 2018a) and another for a more adult audience in December (Naish 2018b). It’s very easy to misunderstand, or ‘mis-frame’, this project as “SCIENTIST WASTES TAX-PAYER MONEY ON LOCH NESS MONSTER NONSENSE” and, indeed, at least one author did make this misunderstanding during the year. What should be noted, I’d argue, is that this is and was a phenomenally successful science outreach programme wherein the Loch Ness Monster was used as the hook to get journalists to cover eDNA and ecological and genetic research, and as such it was absurdly successful (Naish 2018a, b).

Artwork accompanying Naish (2018b). I didn’t write that caption, since the ‘For centuries’ thing is not true. Image: (c)  BBC Focus , Naish (2018b).

Artwork accompanying Naish (2018b). I didn’t write that caption, since the ‘For centuries’ thing is not true. Image: (c) BBC Focus, Naish (2018b).

Late October also saw the release of an episode of the 99% Invisible podcast in which myself, Bob Bakker and John Conway discussed the portrayal of Mesozoic dinosaurs in art. The episode is called Welcome to Jurassic Art and can be found here. Alas, the three of us didn’t really sit around talking (though I have spoken to Bob Bakker, in person, on a few occasions). Instead, our interviews were recorded at different times, at different places. Anyway, it worked out alright.

One of the highlights of the ZSL book-themed event — Jules Howard tells us about the digital duck vagina app. Image: Darren Naish.

One of the highlights of the ZSL book-themed event — Jules Howard tells us about the digital duck vagina app. Image: Darren Naish.

The ZSL zoology books evening. Articles on New Living Animals We Want to Find, Aldrovandi’s monstrous rooster, the second edition of Dinosaurs: How They Lived and Evolved and the pouches of the Sungrebe appeared at TetZoo during November. The article on Aldrovandi’s rooster includes a notable gaff, and an update is due to appear here soon (there is not enough time to do all this stuff, argh!!). Mid-November also saw the Zoological Society of London event From Stoned Sloths to Farting Fish: Comical Tales from the Animal Kingdom, in which myself, Dani Rabaiotti, Jules Howard and Lucy Cooke discussed our adventures in the world of zoology-themed publishing. I mostly spoke about dinosaur sex.

Crew at the public engagement event of December’s Popularising Palaeontology event. L to r: Elsa Panciroli, Mark Witton, Chris Manias, Becky Wragg Sykes, Darren Naish. Image: Darren Naish.

Crew at the public engagement event of December’s Popularising Palaeontology event. L to r: Elsa Panciroli, Mark Witton, Chris Manias, Becky Wragg Sykes, Darren Naish. Image: Darren Naish.

Early in December, I stayed in Bournemouth, UK, for the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation, which has become one of my favourite small meetings. I spoke at last year’s event but not this one. And, later in the month, I attended the Popularising Palaeontology workshop, organised by Chris Manias and hosted by King’s College, London where I listened to some great presentations and participated in the public engagement event.

Herpetology-themed snap cards, obtained at the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation. Image: Darren Naish.

Herpetology-themed snap cards, obtained at the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation. Image: Darren Naish.

And that brings us up to January, a month mostly spent in China and away from the long list of TetZoo-related things I aim to complete in 2019. While in China (specifically, Zigong in Sichuan) I visited Panda Base (properly Chengdu Research Base of Giant Panda Breeding) and the Zigong Dinosaur Museum, among other places.

The entrance to Chengdu Research Base of Giant Panda Breeding, Sichuan, China. Image: Darren Naish.

The entrance to Chengdu Research Base of Giant Panda Breeding, Sichuan, China. Image: Darren Naish.

Thus far, the year’s only TetZoo articles have been those on Heilmann’s Proavis and the life appearance of sauropods. I so wish I could publish more – there is still so much to do. On that note, here’s this…

Achievements for 2018: the pterosaur palaeoneurology paper (Martin-Silverstone et al. 2018), Tetrapod Zoology ver 4 is launched, 2nd edition of Dinosaurs: How They Lived and Evolved sees print (as does Russian translation), Loch Ness magazine articles (Naish 2018a, b), money raised for Eotyrannus monograph, the Dorling Kindersley overhaul, biggest and best TetZooCon so far.

Failures for 2018: not finishing the Eotyrannus monograph, not finishing The Big Book, not making any progress on the Tetrapod Zoology books, Cryptozoologicon Volume 2 or the various dinosaur-themed books planned with John Conway.

Toy thylacines - did you know that there are this many? The grey 3D-printed one - and the painted version at far right (the sitting one) - were obtained at TetZooCon 2018 and were made by Rebecca Groom of palaeoplushies. Image: Darren Naish.

Toy thylacines - did you know that there are this many? The grey 3D-printed one - and the painted version at far right (the sitting one) - were obtained at TetZooCon 2018 and were made by Rebecca Groom of palaeoplushies. Image: Darren Naish.

Ok, so that’s that. How does the year’s blogging do in terms of group representation?

Miscellaneous Musings

Mammals

Lepidosaurs

Pterosaurs

Non-avialan dinosaurs

Birds

Cryptozoology

SpecBio

Tet-Zoo-13th-birthday-Tet-Zoo-13th-birthday-graph-Jan-2018-tiny-Jan-2019-Tetrapod-Zoology-Darren-Naish.jpg

So there we have it. Clearly, 2018 was dominated by non-bird dinosaurs and by articles that can only be classified as generic or miscellaneous and weren’t dedicated to any specific tetrapod group. Amphibians, croc-line archosaurs and so on received no coverage whatsoever. All in all, then, a total and epic failure of what I’ve been trying to achieve here. I figure, by now, that I should stop caring and just aim to write about whatever it is that I write about.

Oh, fish. Image: Darren Naish.

Oh, fish. Image: Darren Naish.

So — happy 13th birthday, Tetrapod Zoology. As ever, I aim to continue to publish as much as I can here: to both present new ideas, new reviews and new takes on zoological topics of interest, and to up-date and correct those things I’ve covered in the past. There is so much to do, and finding time for blogging remains perpetually difficult in view of workload and the never-ending chasing of finances. Plus there’s that textbook I so desperately want to see finished. Here’s your reminder that I’m wholly reliant on funding at patreon, and that I would be able to devote just about all of my work-time to these projects – blogging and the textbook, and the other TetZooniverse-relevant books – if more TetZoo readers were prepared to support me. Huge thanks to those who do so already. Come back soon for another thrilling instalment.

For previous Tet Zoo birthday articles, see...

Refs - -

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

Frigot, R. A. 2018. Pelvic musculature of Vectidraco daisymorrisae and consequences for pterosaur locomotion. In Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications 455, 45-55.

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

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

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

Naish, D. 2018a. Will we ever find Nessie? In Lipscombe-Southwell, A. (ed) BBC Focus Big Book of Mind-Blowing Answers, pp. 16-21.

Naish, D. 2018b. The genetic hunt for Nessie. BBC Focus 329, 66-71.

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

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

Woodward, J. 2018. The Dinosaurs Book. Dorling Kindersley, London.

The Life Appearance of Sauropod Dinosaurs

If you’re a regular TetZoo reader you’ll be familiar with my several articles on the life appearance of Mesozoic dinosaurs, an issue in which I have a special interest. Alas, several of these articles were published at TetZoo ver 2 – the ScienceBlogs years – and hosting issues at the site concerned mean that they’re currently appearing sans all of the many images I so lovingly uploaded. Which is a pain.

A really nice, life-sized model of a titanosaurian sauropod, on display in Romania and constructed by Brian Cooley. Image: Darren Naish.

A really nice, life-sized model of a titanosaurian sauropod, on display in Romania and constructed by Brian Cooley. Image: Darren Naish.

Partly as a consequence, and partly because the issue is on my mind due to several concurrent projects (he says, cryptically), now is a good time to talk once more about the life appearance of Mesozoic dinosaurs, and I’m going to start with sauropods. Sauropods have been covered quite a few times on TetZoo before, as you can see from the list of links at the bottom of this article.

The head, and face especially. We’ll start with the head. Sauropod skulls were proportionally small compared to the overall size of the animals, but not comically so. It’s also worth saying that their eyes – while pretty big in absolute size (based on the size of the eye socket and the sclerotic rings preserved in fossils) – are easy to over-emphasise in illustrations. If you’ve heard that sauropods might have had trunks and thought it reasonable or plausible… well, it’s a poor idea that’s neither reasonable nor plausible, and has a lot counting against it, as covered in the ver 3 article here.

Did sauropods have trunks? NO. It’s the dumbest idea ever, and every piece of evidence counts against it. Several authors have independently explored this idea, including Robert Bakker (upper right) and Bill Munns (lower right). The two images at left are from Knoll  et al .’s (2006) paper refuting the trunk idea. Images: Knoll  et al . (2006),   Bakker (1986)  , Bill Munns ( original here ).

Did sauropods have trunks? NO. It’s the dumbest idea ever, and every piece of evidence counts against it. Several authors have independently explored this idea, including Robert Bakker (upper right) and Bill Munns (lower right). The two images at left are from Knoll et al.’s (2006) paper refuting the trunk idea. Images: Knoll et al. (2006), Bakker (1986), Bill Munns (original here).

Trunks are a no, but did sauropods have ‘lips’ or ‘cheeks’? A whole article could be written on this issue. For now, I’ll summarise things by saying that sauropod skull bone texture indicates that they likely did have extra-oral tissues that mostly covered their teeth (Witton 2018), meaning that they were facially similar to lizards and kin.

A suggestion that some titanosaurs had blade-like cutting sections in the posterior sections of their jaws – so-called ‘guillotine crests’ (Apesteguía 2004) – looks unlikely given that it would require the animals concerned to do their food procurement at the sides of the jaws, rather than at the front like all other sauropods and other dinosaurs. It’s more likely that a mistake has been made here and that the jaw segments thought to support those blade-like, keratinised sections were, while sharp-edged, covered in normal lip tissue. The possibility that diplodocoids and maybe other sauropods too might have had true beaks at the font of the mouth has recently been put forward, but so far only in preliminary fashion.

Apesteguía (2004) argued that some titanosaurs - this is the rhino-sized  Bonitasaura  - had keratinised ‘guillotine crests’ on the edges of the jaws. I think that this is likely a mistake. Image: Apesteguía (2004).

Apesteguía (2004) argued that some titanosaurs - this is the rhino-sized Bonitasaura - had keratinised ‘guillotine crests’ on the edges of the jaws. I think that this is likely a mistake. Image: Apesteguía (2004).

The nostrils and nose. As is now widely known, and near-universally accepted, the external or fleshy nostrils of sauropods were almost certainly not located way up on the forehead, far from the front of the snout, as was long thought based on the retracted position of the bony nostril openings. A more ‘normal’, anterior position for the nostrils was demonstrated by Witmer (2001) who pointed to evidence from blood vessel and nerve impressions and associated cranial hollows, all of which are located on the anterior part of the snout. These indicate the most likely position of the fleshy nostril and associated blood vessel clusters.

The soft dinosaur revolution comes to Sauropod Town. At left, an image by Matt Wedel which shows why we need more soft tissue on our sauropods. At right, Larry Witmer’s (2001) depiction of the different possibilities as goes nostril position in sauropods. (a) is the most likely option based on anatomical data. Image: Mathew Wedel, Witmer (2001).

The soft dinosaur revolution comes to Sauropod Town. At left, an image by Matt Wedel which shows why we need more soft tissue on our sauropods. At right, Larry Witmer’s (2001) depiction of the different possibilities as goes nostril position in sauropods. (a) is the most likely option based on anatomical data. Image: Mathew Wedel, Witmer (2001).

There are also reasons for thinking that the giant, cavernous bony nostril openings and tall nasal bars of some macronarian sauropods supported, and were surrounded by, dome-shaped soft tissue convexities, superficially recalling the bulging nasal regions of some living monitor lizards. Part of my reason for saying this comes from the nasal anatomy of the exquisitely preserved South American titanosaur Sarmientosaurus. Here, an anteriorly projecting bony spine located along the midline and associated bar-like structures on the lateral edges of the large narial fossa – the big bony opening surrounding the bony nostril – indicate that a wide, convex mass of tissue connected the forehead with the sides and front of the snout’s upper surface (Martínez et al. 2016). Sarmientosaurus isn’t unique in this respect but is a particularly good example.

The skulls of some sauropods - this is the titanosaur  Sarmientosaurus  - indicate that there were bulbous nasal structures covering much of the snout region. Image: WitmerLab.

The skulls of some sauropods - this is the titanosaur Sarmientosaurus - indicate that there were bulbous nasal structures covering much of the snout region. Image: WitmerLab.

Indeed, skull openings in general were almost definitely not sunken in appearance or otherwise all that obvious, something that’s being said of archosaurian faces in general as artists and anatomists have learnt to take better attention of the conditions present in living animals (where cranial openings are just about never obvious in the live creature).

Accordingly, sauropod faces were seemingly ‘softer’ and more ‘padded’ than convention would have it, at least some of – perhaps all of – the nasal and forehead region being convex and fleshy, rather than shrink-wrapped and covered in thin skin alone (Witton 2018).

Were sauropod necks mostly semi-horizontal and with a limited range of motion, as argued by Martin (1987) and illustrated at left? Nope; it’s more likely that the necks were often held aloft and far more flexible, as argued by  Taylor  et al . (2009)  and depicted at right. Images: Martin (1987),  Taylor  et al . (2009) .

Were sauropod necks mostly semi-horizontal and with a limited range of motion, as argued by Martin (1987) and illustrated at left? Nope; it’s more likely that the necks were often held aloft and far more flexible, as argued by Taylor et al. (2009) and depicted at right. Images: Martin (1987), Taylor et al. (2009).

The neck. The most remarkable feature of sauropod anatomy is the neck. As goes how the neck was held and how flexible it was, several possibilities have been put forward, some workers arguing that it was held in a mostly horizontal attitude for much of the time and with only a limited range of lateral and vertical flexibility. I’m part of a group who argue for mostly elevated habitual neck poses (even in diplodocids) and a wide range of flexibility (Taylor et al. 2009). Arguments that sauropods must have been horizontal-necked do not take account of the flexibility permitted by cartilage, or – in living animals of all sorts – the ranges of motion that happen at zygapophyseal junctions and at the neck base and head-neck junctions.

There are good reasons for thinking that sauropod necks were habitually inclined upwards, as is typical for terrestrial tetrapods in general, and several fossils are actually preserved with the neck in this orientation. This montage by Greg Paul shows several of the fossil concerned. Image: Paul (1998).

There are good reasons for thinking that sauropod necks were habitually inclined upwards, as is typical for terrestrial tetrapods in general, and several fossils are actually preserved with the neck in this orientation. This montage by Greg Paul shows several of the fossil concerned. Image: Paul (1998).

It's been convention to show the sauropod neck as a featureless tubular structure, like a hose. This is mostly wrong, in part because the shapes of the vertebrae reveal a more unusual cross-sectional shape that varies from one sauropod group to the next. In some sauropods (like mamenchisaurs), the vertebrae are quite narrow and the neck would have looked laterally compressed in places, as it is (for at least some of its length) in giraffes. In diplodocoids – apatosaurines in particular – the neck is narrow close to the head but wide for much of its length, and subtriangular in cross-section, being widest across the neck’s underside. A neck that was quite narrow in its anteriormost quarter or so but was markedly wide for the rest of its length seems to have been the case in brachiosaurs and at least some titanosaurs.

The necks of some sauropods - a good example is the ultra-long-necked  Omeisaurus , photographed at Zigong Dinosaur Museum - are somewhat laterally compressed for at least part of their length. Image: Darren Naish.

The necks of some sauropods - a good example is the ultra-long-necked Omeisaurus, photographed at Zigong Dinosaur Museum - are somewhat laterally compressed for at least part of their length. Image: Darren Naish.

Indeed, the very base of the neck is remarkably broad in some sauropods – Camarasaurus is the classic example, where the neck base is not that different in width from the front of the chest – meaning that the neck would taper gradually along its length if you were looking at the animal from the front or back, or from above or below.

The neck bases of some sauropods - this is the  Camarasaurus  replica on display in London’s NHM - are shockingly broad. It would be wrong to show the neck as a narrow, hose-like object with parallel sides. Image: Darren Naish.

The neck bases of some sauropods - this is the Camarasaurus replica on display in London’s NHM - are shockingly broad. It would be wrong to show the neck as a narrow, hose-like object with parallel sides. Image: Darren Naish.

The vertebrae themselves are very complicated with large hollows on their sides, projecting neural spines on the apices and so on. In the most extreme version of the shrink-wrapping meme promoted by one or two palaeontologists and palaeoartists, sauropod necks have been depicted as if these structures should be visible in the live animal, Ely Kish’s apatosaurines from 1983 being the ultimate example. This was definitely not the case. It remains uncertain how much musculature and soft tissue surrounded the vertebrae, but it was almost certainly (based on the anatomy of living animals) enough to obscure the form of the vertebrae, their bulbous junctions perhaps being visible in sauropods of some or many sorts.

The brilliant, late Ely Kish, one of the best palaeoartists of all time, constructing a scale model of  Apatosaurus  (in preparation for a large painting). Partly on the advice of palaeontologist Dale Russell, she depicted great lateral concavities on the sides of the neck. Image: Russell (1987).

The brilliant, late Ely Kish, one of the best palaeoartists of all time, constructing a scale model of Apatosaurus (in preparation for a large painting). Partly on the advice of palaeontologist Dale Russell, she depicted great lateral concavities on the sides of the neck. Image: Russell (1987).

Necks as display banners. The unprecedented form of the sauropod neck makes it plausible – perhaps even likely – that the neck was used as a display structure. Phil Senter’s suggestion that the size and length of the neck was driven, in evolutionary terms, by its use as a display structure (Senter 2007) is not supported by evidence (Taylor et al. 2011). However, this doesn’t discount co-option of the neck in display, so it really is – while wholly speculative right now – worth taking seriously the possibility that display structures of various kinds could well have adorned sauropod necks. These could include wattles, dewlaps, spiky frills, spines, filaments, inflatable pouches or distensible flags or flaps. A few artists have explored these possibilities, most notably Brian Engh and Emiliano Troco.

As weird as it might seem, it is at least plausible that the sauropod neck was decorated with weird soft-tissue display structures, as depicted here on a diplodocid. Image: Emiliano Troco.

As weird as it might seem, it is at least plausible that the sauropod neck was decorated with weird soft-tissue display structures, as depicted here on a diplodocid. Image: Emiliano Troco.

 Hands. Sauropod hands are extremely odd, and very different from the rounded, elephant-style structures, edged with big nails and/or several claws, shown so often in artistic reconstructions and museum models. Indeed, they’re sufficiently weird and interesting that I’ve written whole articles about them before but, as I said above, these are currently lacking all of their relevant illustrations and are thus all but useless.

Sauropod hands are essentially unique. They’re weird, semi-tubular structures with pillar-like metacarpals. At left, a brachiosaur hand. At right, the hand of the turiasaur  Zby . Images: Anthony Maltese, Darren Naish.

Sauropod hands are essentially unique. They’re weird, semi-tubular structures with pillar-like metacarpals. At left, a brachiosaur hand. At right, the hand of the turiasaur Zby. Images: Anthony Maltese, Darren Naish.

The sauropod hand is essentially a semi-tubular structure formed of elongate metacarpals arranged, pillar-like, in a semi-circle. The posterior surface – corresponding to the palm – was hollow, the consequence being a semilunate area of contact with the ground. We’ve known since at least 1940 that sauropod hands had this very unusual form thanks to fossil tracks (Falkingham et al. 2014), which makes it all the weirder that people have ignored this information and merrily continued giving sauropods elephant-like hands across the decades.

Roland T. Bird’s sauropod track illustrations from the 1940s - shown here - clearly show the true, highly unusual form of the sauropod manus. Shame on those who ignored this information in the following decades. Image:  Falkingham  et al . (2014 ), CC BY 4.0.

Roland T. Bird’s sauropod track illustrations from the 1940s - shown here - clearly show the true, highly unusual form of the sauropod manus. Shame on those who ignored this information in the following decades. Image: Falkingham et al. (2014), CC BY 4.0.

Tracks also show that nails and claws were absent from the hands, except on the thumb where a pointed claw – which varied considerably in exact shape and size from one group to the next – projected inwards and slightly backwards. The thumb claw was lost within Titanosauria, meaning that at least some members of this group lacked nails and claws on their hands altogether. I will concede that poorly defined convexities corresponding to manual digits were present in at least some sauropods, but they still wouldn’t have looked like distinct digits.

Some tracks appear to show that tough semi-conical tubercles projected from the skin on the front and sides of the hand, perhaps giving part of the hand a spiky or tuberculated appearance (Milàn et al. 2005). Maybe these structures had a role in foraging, digging, display or combat, since it’s plausible that they made the hands gnarlier and harder than they would have been otherwise.

Vertical scores associated with sauropod hand prints indicate that at least some of them had tuberculate hand skin, as depicted here (at right) in this modified version of a Greg Paul illustration. Image: Milàn  et al . (2005).

Vertical scores associated with sauropod hand prints indicate that at least some of them had tuberculate hand skin, as depicted here (at right) in this modified version of a Greg Paul illustration. Image: Milàn et al. (2005).

Feet. Three large, curved claws projected anterolaterally from the inner three toes. At least some sauropods possessed four such claws. The outer two toes projected as blunt, rounded convexities in some, most or all sauropods. This is obvious from at least some Brontopodus tracks (Meyer et al. 1994). They might have had nails but were more likely lacking horny structures of any sort and essentially continuous with the rest of the foot’s outer surface. In contrast to the hand, the foot did have a massive fatty pad at its back.

Until recently it was thought that the metatarsals were held at a high angle, the consequence being a short foot shaped like that of an elephant but for the claws (Paul 1987). Data from articulated skeletons and tracks, however, show that the metatarsus was not as erect as argued, in which case the toes were longer and flatter than depicted by some artists.

Tschopp  et al . (2015)   used data from complete  Camarasaurus  hands and feet to produce the skeletal and soft-tissue models you see here (in 1 and 2), and then used these to generate tracks (3). The tracks are an exact match for real fossil tracks. Image: (c)  Tschopp  et al . (2015) .

Tschopp et al. (2015) used data from complete Camarasaurus hands and feet to produce the skeletal and soft-tissue models you see here (in 1 and 2), and then used these to generate tracks (3). The tracks are an exact match for real fossil tracks. Image: (c) Tschopp et al. (2015).

On the subject on feet and limbs, trackways also show that sauropods differed in how they placed their hands and feet, some walking with a very narrow gait (the hands and feet being placed close to the midline), others using a wide gait (where there was some short distance between the hands and feet of the left side versus those of the right), and others being intermediate. It should be remembered that even the widest-gauge sauropods did not walk with their feet all that far apart, but that the legs were almost certainly angled inwards, as is typical in living animals. For more on this issue see Scott Hartman’s article here.

Sauropods were variable in cross-sectional shape and in whether they walked with narrow-gauge or wide-gauge gaits, as depicted here by Scott Hartman. Image:   Scott Hartman’s skeletaldrawing.com

Sauropods were variable in cross-sectional shape and in whether they walked with narrow-gauge or wide-gauge gaits, as depicted here by Scott Hartman. Image: Scott Hartman’s skeletaldrawing.com

The body and tail. Sauropod bodies were variable in cross-sectional shape, length and other details. Diplodocoids, for example, were relatively narrow, deep-bodied and with a tall ridge formed from their neural spines running along the back, while titanosaurs were extremely broad across the hips and must have been just about flat across the back. The thorax may have sloped down ever so slightly in diplodocoids (in part because their forelimbs were shorter than their hindlimbs) whereas the thorax was angled upwards slightly or even markedly in some macronarians, like brachiosaurs and some titanosaurs.

Articulated skeletons show that the tail mostly projected horizontally from the pelvis (a subtle arch at the tail base is present in diplodocoids and some others, in fact), but the macronarians with the upward-sloping bodies also had a downward-sloping tail. A real curiosity is provided by the tails of some mamenchisaurs which seem to have projected upwards at an angle. This has been discussed and illustrated by Paul (2010) and more recently by Hallett & Wedel (2016), but has otherwise gone undiscussed as far as I know. It sounds so odd that surely some mistake has been made… though I really don’t think it has.

Tall neural spines, massive, wing-shaped transverse processes and other structures show that an enormous quantity of musculature would have been obvious along the proximal part of the sauropod tail. This is the tail of … ugh .. Dippy, the cast of  Diplodocus carnegii  until recently on show in London. Image: Darren Naish.

Tall neural spines, massive, wing-shaped transverse processes and other structures show that an enormous quantity of musculature would have been obvious along the proximal part of the sauropod tail. This is the tail of … ugh .. Dippy, the cast of Diplodocus carnegii until recently on show in London. Image: Darren Naish.

As is typical for non-bird dinosaurs, the musculature at the base of the tail was (so we can say from the relevant bony attachment points) evidently enormous and bulky, the proximal part of the tail likely being similar in width to the pelvis and thighs combined.

The integument. We know for definite that sauropods of all groups were scaly animals, since scaly skin impressions and actual preserved skin patches are known for diplodocids, Camarasaurus and titanosaurs at least. A few skin folds here and there were almost certainly present (say, where the limbs met the body, at the limb joints, and at mobile zones in the neck) but a fissured, wrinkled or scored elephant-like skin is a big fat no and every effort should be made to avoid it in artistic depictions.

Haestasaurus  from the English Wealden (shown at left) and a few other sauropods preserve polygonal scales that vary somewhat in size. Image: Darren Naish, Czerkas (1994).

Haestasaurus from the English Wealden (shown at left) and a few other sauropods preserve polygonal scales that vary somewhat in size. Image: Darren Naish, Czerkas (1994).

Polygonal – specifically, pentagonal, hexagonal and heptagonal – scales are known for the macronarian Haestasaurus and some diplodocids, while rounded scales possessing a papilliform texture of tiny bumps are also preserved in a diplodocid specimen (Czerkas 1994). The polygonal structures varied in size somewhat, those present in more mobile parts of the skin (say, the inner crease of the elbow) being smaller than those present in non-mobile sections. Even so, the largest scales were, at most, 60 mm across, which is not large at all on an animal more than 15 m long. This is pretty common in non-bird dinosaurs, by the way. The scales were often so small that they wouldn’t be readily visible from any distance greater than a few metres. The papilliform texture on those diplodocid scales seems to have been widespread across sauropods and means that sauropod skin would have been rough to the touch.

A diplodocid specimen from Wyoming that has scaly skin preserved also preserves tall, conical dermal spines (note: they are not horn-covered bony structures) that must have been arranged along the dorsal midline (Czerkas 1992). These are variable in height (the biggest are 18 cm tall) and seem to have formed a single row along the top of the tail’s proximal part at least. This specimen was originally implied to belong to Diplodocus but is of indeterminate identity. It might belong to Kaatedocus, since remains of this dinosaur come from the exact same quarry.

Large, laterally compressed conical and semi-conical dermal structures lined the upper surface of the tail (at least) in some diplodocids, and perhaps in other diplodocoids and sauropods too. Image: Czerkas (1994).

Large, laterally compressed conical and semi-conical dermal structures lined the upper surface of the tail (at least) in some diplodocids, and perhaps in other diplodocoids and sauropods too. Image: Czerkas (1994).

The conical and semi-conical dermal spines of some diplodocids were variable in height, breadth and shape, as illustrated here. Image: Czerkas (1994).

The conical and semi-conical dermal spines of some diplodocids were variable in height, breadth and shape, as illustrated here. Image: Czerkas (1994).

Were these structures present across all diplodocids, all diplodocoids, all neosauropods or even all sauropods, or were they exclusive to one small clade, perhaps even just to Kaatedocus itself? In the absence of further information we can’t say, but it’s appropriate to depict them on diplodocids and their close kin at least, in the absence of further information. The possibility that shorter and/or taller dermal structures of this sort were present elsewhere on other sauropods exists.

The discovery of those conical and semi-conical spines in a diplodocid led Stephen Czerkas to produce this new look for these dinosaurs. It might be correct, but it is not clear whether the structures were as extensive along the animal’s length as shown here. Image: Czerkas (1992).

The discovery of those conical and semi-conical spines in a diplodocid led Stephen Czerkas to produce this new look for these dinosaurs. It might be correct, but it is not clear whether the structures were as extensive along the animal’s length as shown here. Image: Czerkas (1992).

Horn-covered osteoderms - that is, lumps and nodules with a bony core - were arranged across the backs and flanks of some titanosaurs, specifically the members of the clade Lithostrotia. These structures were variously rounded or oval, sometimes quite flat and sometimes slightly or strongly domed. They appear to have been numerous and prominent in some of the relevant species and would have given their backs and sides a pebbly, armoured appearance.

Colours. There’s a traditional view stating that big dinosaurs were likely grey and plain because big living mammals are, as is the Komodo dragon, I suppose. I’m going to be bold here and say that there’s no reason whatsoever to take any notice of this. Sauropods were not mammals or Komodo dragons. While an argument could be made that camouflage would have been beneficial, and that certain pigments and patterns would have been helpful or necessary for physiological reasons (heat-shedding, heat retention, UV protection and so on), bold and complex patterns and bright, even vibrant colours are all consistent with the ecophysiological demands of being a sauropod, and the good colour vision and complexity of sauropod integument, and likely reliance on visual display, mean that they very likely could have been more like gigantic lizards or birds than elephants. In any case, giraffes, perenties and others show that big, terrestrial animals don’t have to be bland and grey as has often been stated.

The idea that big terrestrial animals have to be bland never was correct. Perenties  Varanus giganteus  might not be that colourful, but they help emphasise the possibility that striking patterns can be present even in very large terrestrial reptiles. Image: (c) Stephen Zozaya.

The idea that big terrestrial animals have to be bland never was correct. Perenties Varanus giganteus might not be that colourful, but they help emphasise the possibility that striking patterns can be present even in very large terrestrial reptiles. Image: (c) Stephen Zozaya.

In the absence of any direct evidence, it seems reasonable to me to reconstruct barred, striped, reticulated or dappled colour schemes, to depict bright colours on faces, necks or other areas considered relevant to display, and to imagine any colours reasonable for big, terrestrial animals associated with woodlands, parks, scrubby places, mangroves and all the other habitats frequented by this long-lived, diverse group of animals.

Want to know more about sauropod life appearance? It sometimes surprises people that there are scarcely any good books dedicated to specific Mesozoic dinosaur groups. For sauropods, the main must-have volume is   Mark Hallett and Matt Wedel’s  The Sauropod Dinosaurs   . It’s really good.    Dinosaurs Past and Present Vol II    is also worth getting, in part because it includes Greg Paul’s (now quite dated, but still useful) article on dinosaur life appearance. Images: amazon ( here  and  here ).

Want to know more about sauropod life appearance? It sometimes surprises people that there are scarcely any good books dedicated to specific Mesozoic dinosaur groups. For sauropods, the main must-have volume is Mark Hallett and Matt Wedel’s The Sauropod Dinosaurs. It’s really good. Dinosaurs Past and Present Vol II is also worth getting, in part because it includes Greg Paul’s (now quite dated, but still useful) article on dinosaur life appearance. Images: amazon (here and here).

If we really want to imagine sauropods as living animals, there is – of course – so much more to say. Body language, posture, gait, social life, anti-predator responses, feeding behaviours, digestive function, sleep and so much more are all things that need to be considered. But that’s where I’ll stop for now. I hope you found this interesting, and we’ll be looking at dinosaur life appearance again sometime soon.

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

For previous TetZoo articles on sauropods, see…

 Refs - -

Apesteguía, S. 2004. Bonitasaura salgadoi gen. et sp. nov.: a beaked sauropod from the Late Cretaceous of Patagonia. Naturwissenschaften 91, 493-497.

Bakker, R. T. 1986. The Dinosaur Heresies. New Theories Unlocking the Mystery of Dinosaurs and their Extinction. William Morrow, New York.

Czerkas, S. A. 1992. Discovery of dermal spines reveals a new look for sauropod dinosaurs. Geology 20, 1068-1070.

Czerkas, S. A. 1994. The history and interpretation of sauropod skin impressions. Gaia 10, 173-182.

Falkingham, P., Bates, K. & Farlow, J. 2014. Historical photogrammetry: Bird’s Paluxy River dinosaur chase sequence digitally reconstructed as it was prior to excavation 70 years ago. PLoS ONE 9, 4: e93247.

Hallett, M. & Wedel, M. J. 2016. The Sauropod Dinosaurs: Life in the Age of Giants. Johns Hopkins University Press, Baltimore.

Knoll, F., Galton, P. M. & López-Antoñanzas, R. 2006. Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus. Geobios 39, 215-221.

Martin, J. 1987. Mobility and feeding of Cetiosaurus (saurischia, sauropoda [sic]) - why the long neck? In Currie, P. J. & Koster, E. H.(eds) Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. Boxtree Books (Drumheller, Alberta), pp. 154-159.

Martínez, R. D. F., Lamanna, M. C., Novas, F. E., Ridgely, R. C., Casal, G. A., Martínez, J. E., Vita, J. R. & Witmer, L. M. 2016. A basal lithostrotian titanosaur (Dinosauria: Sauropoda) with a complete skull: implications for the evolution and paleobiology of Titanosauria. PLoS ONE 11, 4: e0151661. 

Meyer, C. A., Lockley, M. G., Robinson, J. W. & dos Santos, V. F. 1994. A comparison of well-preserved sauropod tracks from the Late Jurassic of Portugal and the western United States: evidence and implications. Gaia 10, 57-64.

Milàn, J., Christiansen, P. & Mateus, O. 2005. A three-dimensionally preserved sauropod manus impression from the Upper Jurassic of Portugal: implications for sauropod manus shape and locomotor mechanics. Kaupia 14, 47-52.

Paul, G. S. 1987. The science and art of restoring the life appearance of dinosaurs and their relatives - a rigorous how-to guide. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present Vol. II. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and London), pp. 4-49.

Paul, G.S. 1998. Terramegathermy and Cope’s rule in the land of titans. Modern Geology 23, 179-217.

Paul, G. S. 2010. Dinosaurs: A Field Guide. A & C Black, London.

Russell, D. A. 1987. Models and paintings of North American dinosaurs. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present, Volume I. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 114-131.

Senter, P. 2007. Necks for sex: sexual selection as an explanation for sauropod dinosaur neck elongation. Journal of Zoology 271, 45-53.

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

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

Tschopp, E., Wings, O., Frauenfelder, T. & Brinkmann, W. 2015. Articulated bone sets of manus and pedes of Camarasaurus (Sauropoda, Dinosauria). Palaeontologia Electronica 18.2.44A: 1-65.

Witmer, L. M. 2001. Nostril position in dinosaurs and other vertebrates and its significance for nasal function. Science 293, 850-853.

Witton, M. P. 2018. The Palaeoartist’s Handbook: Recreating Prehistoric Animals in Art. The Crowood Press, Marlborough.

Heilmann, Thompson, Beebe, Tetrapteryx and the Proavian

Many people familiar with ideas on the early evolution of birds and of bird flight will know of the Proavis or proavian, a hypothetical bird ancestor illustrated and discussed by William Pycraft (1868-1942) in 1906 but made better known by Gerhard Heilmann (1859/1861-1946) during the 1910s and 20s. Versions of this creature were later illustrated in colour by Zdeněk Burian for popular books of the 1960s, 70s and 80s. In the article that you’re about to read, we’re predominantly interested in Heilmann’s take on the Proavis.

Fighting, gliding and climbing proavians, depicted in a cluttered woodland environment, by Gerhard Heilmann. This is typical of Heilmann’s many excellent scenes. He used posed plaster models of the creatures before drawing. This illustration was used in the 1916 Danish version of the book (it first appeared in one of his 1912 articles) but does not appear in the 1926 English version. As we’ll see below, the decision to exclude some images from the 1926 version might have served to make it seem more scholarly than its predecessor. Image: Heilmann (1916).

Fighting, gliding and climbing proavians, depicted in a cluttered woodland environment, by Gerhard Heilmann. This is typical of Heilmann’s many excellent scenes. He used posed plaster models of the creatures before drawing. This illustration was used in the 1916 Danish version of the book (it first appeared in one of his 1912 articles) but does not appear in the 1926 English version. As we’ll see below, the decision to exclude some images from the 1926 version might have served to make it seem more scholarly than its predecessor. Image: Heilmann (1916).

Heilmann was an artist and graphic designer by profession but his interest in bird evolution and anatomy was such that he published a series of articles on the subject for the Danish Ornithological Society between 1912 and 1916 (Nieuwland 2004, Ries 2007).

Self-portrait of Gerhard Heilmann, produced in 1902. Heilmann was an argumentative man often in conflict with “various forms of authority”, and at loggerheads with his own family   (Nieuwland 2004)  . This illustration was included in     Nieuwland (2004)  .

Self-portrait of Gerhard Heilmann, produced in 1902. Heilmann was an argumentative man often in conflict with “various forms of authority”, and at loggerheads with his own family (Nieuwland 2004). This illustration was included in Nieuwland (2004).

Compiled, these articles described Heilmann’s take on the diversity of ancient fossil birds, embryological development in birds and reptiles, and bird anatomy, with the last in the series explaining what these data meant for the appearance and lifestyle of his Proavis, an imagined animal older and anatomically more archaic than Archaeopteryx. These articles were published together in a Danish-language book Vor Nuvaerende Viden om Fuglenes Afstamning (Heilmann 1916), the title meaning Our Current Knowledge of the Descent of Birds.

Heilmann’s The Origin of Birds. This body of work was sufficiently impressive that – while Heilmann was mostly shunned in Denmark – it was well received elsewhere, and he was encouraged by scientists in Germany in particular to translate it into English. The result was the 1926 The Origin of Birds (republished in the USA in 1927), a scholarly and well-illustrated book that was well received internationally and almost immediately deemed the most authoritative work on bird origins. The 1926 volume is quite different from that of 1916 in the illustrations it includes (as we’ll see below), but also in how scientific and speculative the text is, some more fanciful sections on palaeobiology being absent from the English version.

It has been said several times that Heilmann’s work was deemed so impressive that, rather than inspire new work on the subject, it effectively bought research on bird origins to a halt. Most relevant workers (albeit not all) now considered the question of bird origins to be resolved. As noted by historian Ilja Nieuwland, things might not have gone this way had the English-speaking readers of Heilmann’s book known that he was an amateur scientist and an artist by trade (Palm 1997, Nieuwland 2004).

Heilmann’s draftmanship was superb. This illustration (fig. 140 from   Heilmann 1926  ) shows thigh feathering on the chicks of various birds, with 8 being a gliding gecko. “The animal with which we may best compare the bird-ancestor, is the Fringed gecko” [sic] (p. 197). Image:   Heilmann (1926)  .

Heilmann’s draftmanship was superb. This illustration (fig. 140 from Heilmann 1926) shows thigh feathering on the chicks of various birds, with 8 being a gliding gecko. “The animal with which we may best compare the bird-ancestor, is the Fringed gecko” [sic] (p. 197). Image: Heilmann (1926).

Heilmann thought that bird ancestors were likely quadrupedal gliders, similar superficially to living gliding geckos (Heilmann 1926, p. 197). This statement is somewhat odd in view of his endorsement elsewhere of a more cursorial view of these animals, but it seems that he imagined them adopting very different poses when on the ground versus when climbing. Heilmann also argued that birds were not dinosaurs – despite his good understanding of the compelling anatomical similarity between theropod dinosaurs and birds – but were instead the descendants of a group closely related to – and supposedly ancestral to – dinosaurs, termed ‘pseudosuchians’*. This was because of his adherence to ‘Dollo’s Law’ – the (erroneous) idea that a lost anatomical structure cannot be regained – and his mistaken contention that dinosaurs lacked clavicles. If dinosaurs couldn’t be ancestral to birds, the true ancestors must have been another group, hence Heilmann’s use of the more archaic ‘pseudosuchians’, already mooted as possible bird ancestors by Robert Broom in his 1913 description of the South African Euparkeria.

* The term pseudosuchian is today applied to the archosaur lineage that includes crocodylians and all of their extinct relatives. Given the historical baggage that comes with the term, I personally don’t think that this is a good idea at all and would prefer it if another name were used for the lineage concerned (like Crurotarsi)… but that’s an issue for another time.

Euparkeria capensis  has long been intimated as a sort of bird ancestor by those looking for such creatures outside of theropod dinosaurs. Not only is this animal only very distantly related to birds, it’s not even part of the crown-archosaur clade. Image: Taenadoman, CC BY-SA 3.0 ( original here ).

Euparkeria capensis has long been intimated as a sort of bird ancestor by those looking for such creatures outside of theropod dinosaurs. Not only is this animal only very distantly related to birds, it’s not even part of the crown-archosaur clade. Image: Taenadoman, CC BY-SA 3.0 (original here).

By combining features common to Archaeopteryx as well as to the ‘pseudosuchians’ Aetosaurus, Euparkeria, Ornithosuchus and Saltoposuchus, Heilmann (1916, 1926) described how he invented a creature that looked somewhat like a theropod but had a more archaic skull and foot, retained a fourth metacarpal, and had a smaller pelvis with far shorter pubic and ischial bones.

Heilmann’s skeletal reconstruction of Proavis, as depicted in the 1916 Danish version of the book… but not in the 1926 English version. Note the lack of long feathers on the hindlimbs. Incidentally, note also that Heilmann was partly responsible for encouraging the belief that forelimb feathers did not cover the hands during the earliest stages of bird evolution. Image: Heilmann (1916).

Heilmann’s skeletal reconstruction of Proavis, as depicted in the 1916 Danish version of the book… but not in the 1926 English version. Note the lack of long feathers on the hindlimbs. Incidentally, note also that Heilmann was partly responsible for encouraging the belief that forelimb feathers did not cover the hands during the earliest stages of bird evolution. Image: Heilmann (1916).

Heilmann, Thompson and D’Arcy Thompson grids. It’s a matter of some interest that Heilmann didn’t just guess what the imagined morphology of the proavian would be, as might be assumed given the English edition of his book. Instead, he used a technique whereby the relevant anatomical regions were mapped on to a grid and then distorted to mimic the evolutionary process. This grid-based deformation process was pioneered by D’Arcy Wentworth Thompson (1860-1948) and was explained most thoroughly in his 1917 book On Growth and Form (Thompson 1917). It was actually put forward beforehand in 1915 (Thompson 1915).

Thompson applied his Cartesian grid technique to many animal lineages, his aim being to show that mathematically predictable transformation in one or more anatomical regions could result in profound anatomical change. Here’s one of my favourite examples: how you can derive a molid sunfish ( Mola  was known to Thompson as  Orthagoriscus ) from a porcupinefish. Image: Thompson (1917).

Thompson applied his Cartesian grid technique to many animal lineages, his aim being to show that mathematically predictable transformation in one or more anatomical regions could result in profound anatomical change. Here’s one of my favourite examples: how you can derive a molid sunfish (Mola was known to Thompson as Orthagoriscus) from a porcupinefish. Image: Thompson (1917).

By placing grids on top of diagrams of related animals (say, the skull of a human and a chimp), Thompson showed how the skewing of the grid in a certain direction “would result in the sort of changes that would allow the emergence of a new species” (Naish 2017, p. 116). These grids are generally termed D’Arcy Thompson transformation grids, Cartesian transformations or Cartesian grids, and their use was quite popular in the evolutionary literature of the early 20th century. Heilmann was inspired to use the technique after Thompson wrote to him about bird evolution in 1915, and the two wrote to each other on many occasions about the technique and its application to ideas on the evolution of horses, hominids and birds. They also discussed how the resulting illustrations could be used in Thompson’s On Growth and Form (Ries 2007).

Heilmann included these Cartesian transformations in the 1916 version of his book, but they weren’t included in the 1926 English version. His take on Proavis was not, therefore, simple guesswork. Image: this montage is from Witmer (1991) but the originals are from Heilmann (1916).

Heilmann included these Cartesian transformations in the 1916 version of his book, but they weren’t included in the 1926 English version. His take on Proavis was not, therefore, simple guesswork. Image: this montage is from Witmer (1991) but the originals are from Heilmann (1916).

Heilmann’s commitment to the technique is demonstrated by the fact that the 1916 Danish edition of his book includes grids that depict the inferred evolution of the avian skull, forelimb and pelvis (I don’t own a copy of the Danish edition, but the diagrams are included in both Witmer (1991) and Ries (2007)). With Ornithosuchus and Euparkeria as ‘starting points’ and Archaeopteryx and modern birds as ‘end points’*, Heilmann used the grids to create relevant intermediates, the results allowing him to generate, piecemeal, his Proavis (Heilmann 1916). A lateral view of the hypothetical animal – showing it in a neural, non-dynamic walking pose – was included in the Danish edition (Heilmann 1916, Ries 2007).

* It should be noted that Heilmann’s Archaeopteryx was not accurate, some of its details (in the skull especially) being semi-hypothetical and informed by Heilmann’s interpretation of ‘pseudosuchians’ like Aetosaurus.

The more dynamic, climbing version of Heilmann’s Proavis skeletal reconstruction, the only version of the reconstruction included in the 1926 English version of his book. Image:   Heilmann (1926)  .

The more dynamic, climbing version of Heilmann’s Proavis skeletal reconstruction, the only version of the reconstruction included in the 1926 English version of his book. Image: Heilmann (1926).

Remarkably, none of this was included in the English edition at all, this creating the impression that Heilmann simply invented the proavian via intuition and guesswork. He didn’t: it was generated via a testable, repeatable method (albeit using flawed anatomical data). Furthermore, the fact that Heilmann’s skeletal reconstruction of Proavis included in the English edition (Heilmann 1926) is the more dynamic climbing version of the creature gives it a less scientific, more speculative, artistic air than the walking version of 1916.

Heilmann, Beebe and Tetrapteryx. When it came to feathering and other aspects of the integument, Heilmann noted his thoughts on another Proavis-type animal, this time the one invented by Charles William Beebe in 1915. Beebe (1877-1962) was an interesting person. He was a naturalist, ecologist, ornithologist, marine biologist, author and explorer, among other things. Today, he might be best known for the deep-water observations he made while in a bathysphere off the coast of Nonsuch Island off Bermuda during the 1930s.

William Beebe’s hypothetical tetrapteryx creature. A pre- Archaepteryx  glider. You’ll note that Beebe was a pretty good artist [UPDATE: this is an error. The illustrations in this publication were by Dwight Franklin, not by Beebe!]. Image: Beebe (1915).

William Beebe’s hypothetical tetrapteryx creature. A pre-Archaepteryx glider. You’ll note that Beebe was a pretty good artist [UPDATE: this is an error. The illustrations in this publication were by Dwight Franklin, not by Beebe!]. Image: Beebe (1915).

When it came to bird origins, Beebe thought that birds originated via a ‘tetrapteryx’ phase where large feathers on the proximal hindlimb were present and functioned in slowing descent during leaping and gliding (Beebe 1915). Both the fossils of Archaeopteryx and the nestlings of living bird species demonstrated the antiquity of these ‘pelvic wings’, Beebe (1915) said. Heilmann (1926) disagreed. His own examination of bird nestlings pertaining to bird lineages across the family tree resulted in his “complete disappointment, for what I found was wholly negative; there was not in any of them the slightest trace of a “pelvic wing”” (p. 194).

As for Archaeopteryx, Heilmann (1926) argued that the hindlimbs were arranged such that they could never have had an aerodynamic function, nor was there evidence for long hindlimb feathering of the sort consistent with ‘pelvic wings’. Ergo, Heilmann illustrated Archaeopteryx without long hindlimb feathering, and he didn’t think that Proavis had them either: his skeletal reconstructions (both the 1916 walking version and 1926 climbing one) lack them (the 1926 climber has short hindlimb feathers, similar in length to those elsewhere on the body) and the gliding proavians included in his various drawings clearly lack them as well. His most famous proavian figure – the semi-erect one standing in a conifer tree (shown below) – does have long feathers on the rear edge of the thigh, though they certainly aren’t as long as they should be as per Beebe’s tetrapteryx model.

The iconic tree-dwelling proavian. The iconic tree-dwelling proavian I have in mind here is one of Heilmann’s best-known and most frequently reproduced illustrations. It has what appears to be great symbolic significance if you’ve read the English version of The Origin of Birds since it’s the very last image in the entire book (Heilmann 1926, p. 199), thus appearing as a visual metaphor for an imagined phrase: “Finally, I can reconstruct the real ancestor of birds, and it looked like this”.

Heilmann’s most iconic version of his Proavis. Note the long feathers on the back of the thigh and how they’re invisible and thus not deemed important in the gliding individual in the background. Heilmann said that the long feathers close to the base of the tail would produce the required lift in this region. Image:   Heilmann (1926)  .

Heilmann’s most iconic version of his Proavis. Note the long feathers on the back of the thigh and how they’re invisible and thus not deemed important in the gliding individual in the background. Heilmann said that the long feathers close to the base of the tail would produce the required lift in this region. Image: Heilmann (1926).

I was surprised to learn that Beebe produced what looks like his own version of this illustration, and inserted it as a plate in a book published in 1915. However, the image itself is not dated 1915 and does not definitely pre-date Heilmann’s illustration. My suspicion – and that of Paul Stewart, who kindly brought my attention to the image – is that Beebe copied Heilmann’s proavian for his own personal use (the illustration was not intended for publication*). This in itself is interesting since it could mean that Beebe personally endorsed Heilmann’s view of Proavis and regarded it as a ‘descendant’ of his tetrapteryx creature… which it wasn’t, given that Heilmann rejected Beebe’s idea, as we saw earlier. Again, my thanks to Paul Stewart for sharing this image with me and permitting its use here.

* Thanks to Jonathan Kane, I’ve learnt that it was previously published in Tim Berra’s 1977 William Beebe: An Annotated Bibliography.

William Beebe seems to have produced his own take on Heilmann’s iconic proavian scene. We’re presuming that it was produced some time after Heilmann’s illustration was, but this hasn’t yet been fully confirmed. The illustration appears here courtesy of Paul Stewart.

William Beebe seems to have produced his own take on Heilmann’s iconic proavian scene. We’re presuming that it was produced some time after Heilmann’s illustration was, but this hasn’t yet been fully confirmed. The illustration appears here courtesy of Paul Stewart.

Today, the significance of Beebe’s tetrapteryx idea is a bit uncertain and, frankly, depends on who you ask. Archaeopteryx probably did have long feathers on the hindlimbs (Longrich 2006), and they perhaps had an aerodynamic role if this animal indulged in aerial locomotion (which it likely did). Furthermore, the discovery of prominent ‘hindlimb wings’ in other feathered dinosaurs (most notably the Chinese dromaeosaur Microraptor) has led some experts to note the predictive power of Beebe’s suggestion (Kane et al. 2016) and to favour the presence and importance of hindlimb feathers in the earliest birds (Longrich 2006, Chatterjee & Templin 2007, Zheng et al. 2013, Xu et al. 2014). However, it’s not at all certain that large, aerodynamically ‘functional’ hindlimb feathers were present consistently across the lineages concerned or that they had the role that Beebe imagined (O’Connor & Chang 2015).

Recent work shows that  Archaeopteryx  really did have long feathers on the proximal portions of its hindlimbs (these reconstructions are by Longrich (2006)), though they weren’t as long as the feathers Beebe imagined for his tetrapteryx stage creature. Image: Longrich (2006).

Recent work shows that Archaeopteryx really did have long feathers on the proximal portions of its hindlimbs (these reconstructions are by Longrich (2006)), though they weren’t as long as the feathers Beebe imagined for his tetrapteryx stage creature. Image: Longrich (2006).

As always, there’s more to say and this article is already longer than intended. I also wanted to talk about Pycraft’s initial concept of the Proavis, and also Burian’s depiction, since he didn’t simply copy Heilmann’s illustrations but actually produced yet another hypothetical creature. These things will have to wait to another time. Also worthy of further discussion is the point made earlier about Heilmann’s influence happening despite his status as a ‘mere’ artist, a theme that has parallels elsewhere in the history of vertebrate palaeontology and is relevant to my writings on meme perpetuation, the scientific acceptance of feathering in non-bird dinosaurs and much else besides. I aim to explore these topics (and others) in future articles.

Before Heilmann, there was Pycraft’s proavian of 1906. We’ll have to discuss this creature and its backstory another time. Image:   Pycraft (1910)  .

Before Heilmann, there was Pycraft’s proavian of 1906. We’ll have to discuss this creature and its backstory another time. Image: Pycraft (1910).

On which note, please consider supporting this blog at patreon if you don’t already do so. The more support I receive, the more time I can spend generating new content. Thanks so much.

For previous TetZoo articles relevant to the issues covered here, see…

Refs - -

Beebe, C. W. 1915. A tetrapteryx phase in the ancestry of birds. Zoologica 2, 38-52.

Chatterjee, S. & Templin, R. J. 2007. Biplane wing planform and flight performance of the feathered dinosaur Microraptor gui. Proceedings of the National Academy of Sciences 104, 1576-1580.

Heilmann, G. 1916. Vor Nuvaerende Viden om Fuglenes Afstamning. Unknown publisher, Copenhagen.

Heilmann, G. 1926. The Origin of Birds. Witherby, London.

Kane, J., Willoughby, E. & Keesey, T. M. 2016. God’s Word or Human Reason? An Inside Perspective on Creationism. Inkwater Press.

Longrich, N. 2006. Structure and function of hindlimb feathers in Archaeopteryx lithographica. Paleobiology 32, 417-431.

Naish, D. 2017. Evolution in Minutes. Quercus, London.

Nieuwland, I. J. J. 2004. Gerhard Heilmann and the artist’s eye in science, 1912-1927. www.PalArch.nl., vertebrate palaeontology 3, 2.

O’Connor, J. & Chang, H. 2015. Hindlimb feathers in paravians: primarily “wings” or ornaments? Biology Bulletin 42, 616-621.

Palm, S. 1997. The Origin of Flapping Flight in Birds. Svend Plam, Ballerop.

Pycraft, W. P. 1910. A History of Birds. Methuen & Co, London.

Ries, C. J. 2007. Creating the Proavis: bird origins in the art and science of Gerhard Heilmann 1913-1926. Archives of Natural History 34, 1-19.

Thompson, D. W. 1915. Morphology and mathematics. Transactions of the Royal Society of Edinburgh 50, 857-895.

Thompson, D. W. 1917. On Growth and Form. Cambridge University Press, Cambridge.

Witmer, L. M. 1991. Perspectives on avian origins. In Schultze, H.-P. & Trueb, L. (eds) Origins of the Higher Groups of Tetrapods: Controversy and Consensus. Cornel University Press (Ithaca, London), pp. 427-466.

Xu, X., Zhou, Z., Dudley, R., Mackem, S., Chuong, C.-M., Erickson, G. M. & Varricchio, D. J. 2014. An integrative approach to understanding bird origins. Science 346 (6215), 1253293.

Zheng, X., Zhou, Z., Wang, X., Zhang, F., Zhang, X., Wang, Y., Wei, G., Wang, S. & Xu, X. 2013. Hind wings in basal birds and the evolution of leg feathers. Science 339, 1309-1312.

The Most Amazing TetZoo-Themed Discoveries of 2018

As we hurtle toward the end of the year – always a scary thing because you realise how much you didn’t get done in the year that’s passed – it’s time to look back at just a little of what happened in 2018. This article is not anything like a TetZoo review of 2018 (I’ll aim to produce something along those lines in early 2019), but, rather, a quick look at some of the year’s neatest and most exciting zoological (well, tetrapodological) discoveries. As per usual, I intended to write a whole lot more – there are so many things worthy of coverage – and what we have here is very much an abridged version of what I planned.

Animals we will meet below, a montage. Images: (c) Philippe Verbelen, (c) Kristen Grace, Florida Museum of Natural History,  Graham  et al . (2018) , CC BY-SA 4.0.

Animals we will meet below, a montage. Images: (c) Philippe Verbelen, (c) Kristen Grace, Florida Museum of Natural History, Graham et al. (2018), CC BY-SA 4.0.

Thanks as always to those supporting me at patreon. Time is the great constraint (and finance, of course), and the more support I have, the more time I can spend on producing blog content. Anyway, to business…

The Rote leaf warbler. New passerine bird species are still discovered on a fairly regular basis; in fact three were named in 2018*. One of these is especially remarkable. It’s a leaf warbler, or phylloscopid, endemic to Rote in the Lesser Sundas, and like most members of the group is a canopy-dwelling, insectivorous, greenish bird that gleans for prey among foliage. Leaf warblers are generally samey in profile and bill shape, so the big deal about the new Rote species – the Rote leaf warbler Phylloscopus rotiensis – is that its bill is proportionally long and curved, giving it a unique look within the group. It superficially recalls a tailorbird. Indeed, I think it’s likely that the species would be considered ‘distinct enough’ for its own genus if there weren’t compelling molecular data that nests it deeply within Phylloscopus (Ng et al. 2018).

* The others are the Cordillera Azul antbird Myrmoderus eowilsoni and the Western square-tailed drongo Dicrurus occidentalis.

A Common chiffchaff  Phylloscopus collybita  encountered in western Europe, a familiar Eurasian-African phylloscopid leaf warbler. Image: Darren Naish.

A Common chiffchaff Phylloscopus collybita encountered in western Europe, a familiar Eurasian-African phylloscopid leaf warbler. Image: Darren Naish.

The story of the Rote leaf warbler’s discovery is interesting in that it’s yet another recently discovered species whose existence and novelty was suspected for a while. Colin Trainor reported leaf warblers on Rote in 2004 but never got a good look at them, Philippe Verbelen observed them in 2009 and realised how anatomically unusual they were, and it wasn’t until 2015 that a holotype specimen was procured (Ng et al. 2018). I’ve mentioned before the fact that documenting and eventually publishing a new species is rarely an instant see it catch it publish it event, but a drawn-out one that can take decades, and here we are again. Also worth noting is that the existence of a leaf warbler on Rote was not predicted based on our prior knowledge of leaf warbler distribution in view of the deep marine channel separating Rote from Timor and lack of any prior terrestrial connection. Yeah, birds can fly, but members of many groups prefer not to cross deep water channels. In this case, this did, however, happen and most likely at some point late in the Pliocene (Ng et al. 2018).

Rote leaf warbler in life, a novel member of an otherwise conservative group. Image: (c) Philippe Verbelen.

Rote leaf warbler in life, a novel member of an otherwise conservative group. Image: (c) Philippe Verbelen.

Rote has yielded other new passerines in recent years – the Rote myzomela Myzomela irianawidodoae (a honeyeater) was named in 2017 – and it’s possible that one or two others might still await discovery there.

Neanderthal cave art. Hominins don’t get covered much at TetZoo, which is weird given the amazing pace of relevant recent discoveries and the fact that they’re totally part of the remit. I mostly don’t cover them because I feel they’re sufficiently written about elsewhere in the science blogging universe, plus I tend to be preoccupied with other things. Nevertheless, I take notice, and of the many very interesting things published in 2018 was Hoffman et al.’s (2018) announcement of several different pieces of Spanish rock art, seemingly made by Neanderthals Homo neanderthalensis. The art concerned involves hand stencils, abstract lines, squares and amorphous patches of pigment, always marked in red.

Red abstract markings, discovered in several Spanish caves, are old, and in fact were seemingly made by hominins long before  H. sapiens  moved into Europe. The red sinuous marking and system of squares and lines near the middle of this photo are purported to have been made by Neanderthals (other images, depicting animals and present adjacent to these markings, were seemingly created more recently by  H. sapiens  individuals). Image: (c) P. Saura.

Red abstract markings, discovered in several Spanish caves, are old, and in fact were seemingly made by hominins long before H. sapiens moved into Europe. The red sinuous marking and system of squares and lines near the middle of this photo are purported to have been made by Neanderthals (other images, depicting animals and present adjacent to these markings, were seemingly created more recently by H. sapiens individuals). Image: (c) P. Saura.

The main reason for the attribution of this art to Neanderthals is its age: uranium-thorium dating shows that it’s older than 64ka, which therefore makes it more than 20ka older than the time at which H. sapiens arrived in Europe (Hoffman et al. 2018). That seems compelling, and it’s consistent with a building quantity of evidence for Neanderthal cultural complexity which involves the use of shells, pigments, broken stalagmites and so on.

One of the most famous pieces of claimed Neanderthal rock art: the Gorham's Cave ‘hashtag’ from Gibraltar. Image: (c) Stewart Finlayson.

One of the most famous pieces of claimed Neanderthal rock art: the Gorham's Cave ‘hashtag’ from Gibraltar. Image: (c) Stewart Finlayson.

I should add here, however, that I’m slightly sceptical of the use of age as a guide to species-level identification. Why? Well, we have evidence from elsewhere in the fossil record that the range of a hominin species can be extended by around 100ka without serious issue (witness the 2017 announcement of H. sapiens remains from north Africa; a discovery which substantially increased the longevity of our species). In view of this, would a 20ka extension of H. sapiens’ presence in Europe be absolutely out of the question? Such a possibility is not supported by evidence yet, and I don’t mean to appear at all biased against Neanderthals.

A tiny Cretaceous anguimorph in amber, and other Mesozoic amber animals. As you’ll know if you follow fossil-themed news, recent years have seen the discovery of an impressive number of vertebrate fossils in Cretaceous amber, virtually all of which are from Myanmar and date to around 99 million years old. They include tiny enantiornithine birds, various feathers (most recently racquet-like ‘rachis dominated feathers’), the tiny snake Xiaophis, early members of the gecko and chameleon lineages and the small frog Electrorana. Many of these finds were published in 2018 and any one could count as an ‘amazing’ discovery.

The  Barlochersaurus winhtini  holotype, from Daza  et al . (2018).

The Barlochersaurus winhtini holotype, from Daza et al. (2018).

However, there’s one fossil in particular that I find ‘amazing’, and it hasn’t received all that much coverage. It’s the tiny (SVL* 19.1 mm!), slim-bodied anguimorph Barlochersaurus winhtini, named for a single, near-complete specimen subjected to CT-scanning (Daza et al. 2018). Remarkable images of its anatomical details are included in Daza et al.’s (2018) paper. It has short limbs, pentadactyl hands and feet and a slim, shallow, bullet-shaped skull. Phylogenetic study finds it to be somewhere close to, or within, Platynota (the clade that includes gila monsters and kin, and monitors and kin), or perhaps a shinisaurian (Daza et al. 2018). It could be a specialised dwarf form, or somehow more reflective of the ancestral bauplan for these anguimorph groups. Either way, it’s exciting and interesting. What next from Burmese amber?

* snout to vent length

Barlochersaurus  in life. It’s about the size of a paperclip. Image: (c) Kristen Grace, Florida Museum of Natural History ( original here ).

Barlochersaurus in life. It’s about the size of a paperclip. Image: (c) Kristen Grace, Florida Museum of Natural History (original here).

The Reticulated Siren. Sirens are very special, long-bodied aquatic salamanders with reduced limbs and bushy external gills. They’re very weird. They can reach 95 cm in length (and some fossil species were even larger), lack hindlimbs and a pelvis, have a horny beak and pavements of crushing teeth, and eat plants in addition to gastropods, bivalves and other animal prey. A longish article on siren biology and evolution can be found here at TetZoo ver 3.

A life reconstruction of the Cretaceous siren  Habrosaurus , showing features typical of the group. This animal could reach 1.5 m in total length. Image: Darren Naish (prepared for my in-prep texbook The Vertebrate Fossil Record,   on which go here  ).

A life reconstruction of the Cretaceous siren Habrosaurus, showing features typical of the group. This animal could reach 1.5 m in total length. Image: Darren Naish (prepared for my in-prep texbook The Vertebrate Fossil Record, on which go here).

Until recently, just four living siren species were recognised. But it turns out that indications of a fifth – endemic to southern Alabama and the Florida panhandle – have been around since 1970 at least. Furthermore, they pertain to a big species, similar in size to the Great siren Siren lacertina. Known locally as the ‘leopard eel’ (a less than ideal moniker, given that there’s a real eel that already goes by this name), this animal has been published by Sean Graham and colleagues in the open-access journal PLoS ONE (Graham et al. 2018) wherein it’s formally christened the Reticulated siren S. reticulata. It reaches 60 cm in total length, has dark spots across its dorsal surface and a proportionally smaller head and longer tail than other Siren species.

A museum specimen of the species has been known since 1970 when its finder noted that it did “not conform” to descriptions of known species, and live specimens were collected by David Steen and colleagues in 2009 and 2014. Again, note that discovery and recognition was a drawn-out process. The discovery has, quite rightly, received a substantial amount of media coverage, and many interesting articles about the find are already online. Many of you will already know of David Steen due to his social media presence and Alongside Wild charity (which I’m proud to say I support via pledges at patreon).

The Reticulated siren paratype specimen, as described by  Graham  et al . (2018) . Image:  Graham  et al . (2018) , CC BY-SA 4.0.   Original here.

The Reticulated siren paratype specimen, as described by Graham et al. (2018). Image: Graham et al. (2018), CC BY-SA 4.0. Original here.

The idea that a new living amphibian species 60 cm long might be discovered anew in North America in 2018 is pretty radical. I’m reminded of the 2009 TetZoo ver 2 article ‘The USA is still yielding lots of new extant tetrapod species’ (which is less fun to look at than it should be, since images aren’t currently showing at ver 2). Furthermore, Graham et al. (2018) discovered during their molecular phylogenetic work that some other siren species are not monophyletic but likely species complexes, in which case taxonomic revision is required and more new species will probably be named down the line.

And that’s where I must end things, even though there are easily another ten discoveries I’d like to write about. This is very likely the last article I’ll have time to deal with before Christmas. As I write, I’m preparing to leave for the Popularising Palaeontology conference which happens in London this week (more info here), and then there are Christmas parties and a ton of consultancy jobs to get done before the New Year. On that note, I’ll sign off with a festive message, as is tradition. Best wishes for the season, and here’s to a fruitful and action-packed 2019. Special thanks once again to those helping me out at patreon.

TetZooniverse-Christmas-2018-tiny-from-Darren-Naish.jpg

For previous TetZoo articles relevant to various of the subjects covered here, see…

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.

Graham, S. P., Kline, R., Steen, D. A. & Kelehear, C. 2018. Description of an extant salamander from the Gulf Coastal Plain of North America: the Reticulated Siren, Siren reticulata. PLoS ONE 13 (12): e0207460.

Hoffman, D. L., Standish, C. D., García-Diez, M., Pettitt, P. B., Milton, J. A., Zilhão, J., Alcolea-González, J. J., Cantelejo-Duarte, P., Collado, H., de Balbín, R., Lorblanchet, M., Ramos-Muñoz, J., Weniger, G.-Ch. & Pike, A. W. G. 2018. U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art. Science 359, 912-915.

Ng, N. S. R., Prawiradilaga, D. M., Ng, E. Y. X., Suparno, Ashari, H., Trainor, C., Verbelen, P. & Rheindt, F. E. 2018. A striking new species of leaf warbler from the Lesser Sundas as uncovered through morphology and genomics. Scientific Reports 8: 15646.

Up Close and Personal With the Crystal Palace Dinosaurs

Like many of us, I have long enjoyed looking at the Crystal Palace dinosaurs and other prehistoric animal models, created in 1854, still on show more than 160 years later, and providing a remarkable showcase of ancient life as it was imagined at the time. But I’ve only ever seen them from afar. How fantastic would it be to examine them up close? Well…

You’ve seen the Crystal Palace dinosaurs before (or images of them, anyway), but you might not have seen them up-close like this. Neither had I prior to this very special visit. Image: Darren Naish.

You’ve seen the Crystal Palace dinosaurs before (or images of them, anyway), but you might not have seen them up-close like this. Neither had I prior to this very special visit. Image: Darren Naish.

Way back in September 2018, I was fortunate enough to attend the Crystal Palace Dinosaur Days event, part of the Heritage Open Days weekend occurring across the UK on the weekend concerned. I gave a talk and also led a tour around the prehistoric animal models (focusing on the reptiles and amphibians alone). Adrian Lister (of mammoth and Megaloceros fame) led a tour too, Mark Witton gave a talk on ‘Palaeoart After Crystal Palace’, and much else happened besides. I also have to mention the 3D-printed models of the dinosaurs made by Perri Wheeler. How I would love for these to be commercially available: I’m sure they’d be a success. So, it was a great event; well done Ellinor Michel and everyone else involved in the Friends of Crystal Palace Dinosaurs group (follow them on Twitter at @cpdinosaurs) for putting it together.

Perri Wheeler’s brilliant models of the three Crystal Palace dinosaurs (from back to front:  Megalosaurus ,  Hylaeosaurus ,  Iguanodon ). As a pathological collector of model dinosaurs, I sure would like to own a set - but I also sure would like for these models (or a set very similar to them) to be commercially available. Image: Darren Naish.

Perri Wheeler’s brilliant models of the three Crystal Palace dinosaurs (from back to front: Megalosaurus, Hylaeosaurus, Iguanodon). As a pathological collector of model dinosaurs, I sure would like to own a set - but I also sure would like for these models (or a set very similar to them) to be commercially available. Image: Darren Naish.

The real thrill, however, was not the talks nor the presence of the amazing and sometimes spectacularly good speakers but the fact that we were awarded special, up-close access to the prehistoric animal models. A dream come true. As you’ll know if you’ve visited Crystal Palace or read about it, the models are located on islands surrounded by a snaking waterway. In other words, they aren’t readily accessible. For the duration of Dinosaurs Days, however, a temporary bridge had been erected and – like Lord Roxton striding across a felled tree to Maple White Land – we made the crossing and stepped into a bygone era.