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.

New Living Animals We Want to Find

As a regular denizen of the TetZooniverse, you may well remember the July 2017 article ‘Fossils We Want to Find’ in which I discussed a list of hypothetical fossil things that we might one day discover but haven’t yet. Wouldn’t it be fun to do the same sort of thing with extant species; that is, with discoveries pertaining to living, breathing animals? Over at the Zoology for Enthusiasts facebook group (a spinoff of the Tetrapod Zoology facebook group), Jordan Fryer suggested doing exactly this, and consequently people have been coming up with their own suggested living animals that might await discovery. Because this seemed like a lot of fun (and a chance to discuss some really neat and unusual stuff), I thought I’d give it a go.

The 2017 precursor to the article you’re reading here was all about fossil animals. It included this photo, which shows me in the act of discovering a dinosaur bone in the Moroccan Sahara. Image: Richard Hing.

The 2017 precursor to the article you’re reading here was all about fossil animals. It included this photo, which shows me in the act of discovering a dinosaur bone in the Moroccan Sahara. Image: Richard Hing.

Naturally, any list of this sort is horribly subjective, reflecting the interests and biases of the person compiling the list, but so be it. It also seems all too easy to turn any compilation into a ‘list of most discoverable cryptids’: for those of you who don’t know, I have a long-standing interest in cryptozoology and have published on it quite frequently (see Naish (2017) for starters). For the most part, I’ve not done this, though read on.

Many of my thoughts on mystery animals can be found in my   2017 book  Hunting Monsters   . I am not - sorry - much impressed by the case for such supposed animals as the mokele-mbembe, an artistic reconstruction of which is shown at right. Image: David Miller, in   Mackal (1987)  .

Many of my thoughts on mystery animals can be found in my 2017 book Hunting Monsters. I am not - sorry - much impressed by the case for such supposed animals as the mokele-mbembe, an artistic reconstruction of which is shown at right. Image: David Miller, in Mackal (1987).

I’ve also mostly excluded hypothetical discoveries that are inspired by the creatures of cryptozoology but could arguably be considered independent of the cryptozoological literature. In part this is because I don’t think they’re plausible or worth considering, but it’s also because they’re cliched and the opposite of original. So, no ‘living sauropods from the Congo’ or ‘living plesiosaurs in Loch Ness’, for example.

As for what I have selected: well, some of my suggestions are sillier than others, and some are perhaps not that interesting to non-specialists. But, whatever. Feel free to dissect my suggestions in the comments, and perhaps come up with your own.

Among my suggested ‘fossils we want to find’ are protobats (like the hypothetical examples shown at left, from Graham (2002)) and a good skeleton of the giant hominid  Gigantopithecus blacki . This ilustration of a lower jaw is from Simons & Ettel's (1970) magazine article. Images: Graham (2002), Simons & Ettel (1970).

Among my suggested ‘fossils we want to find’ are protobats (like the hypothetical examples shown at left, from Graham (2002)) and a good skeleton of the giant hominid Gigantopithecus blacki. This ilustration of a lower jaw is from Simons & Ettel's (1970) magazine article. Images: Graham (2002), Simons & Ettel (1970).

A habitually bipedal, large, non-human hominid. Whatever you think of all those stories, anecdotes and sightings about bigfoot, yeti, almas, orang-pendek, yowie and so on and on, the fact remains that the discovery of a large, bipedal non-human hominid – whether it be a pongine, hominine, or member of another hominid lineage – would be a huge deal. It would not just be one of the most newsworthy creatures to ever be discovered; it would also have enormous ramifications for our understanding of hominid evolution and potentially the human condition itself.

Are crypto-hominids a cultural phenomenon more than a zoological one? I’ve argued for both possibilities at different times. Whatever… for the purposes of the article you’re reading now, I hope we can agree that the discovery of such an animal would be high on any hypothetical ‘wants’ list. Image: Darren Naish.

Are crypto-hominids a cultural phenomenon more than a zoological one? I’ve argued for both possibilities at different times. Whatever… for the purposes of the article you’re reading now, I hope we can agree that the discovery of such an animal would be high on any hypothetical ‘wants’ list. Image: Darren Naish.

It would also – if relating to North America or northern Eurasia in particular – very likely have a significant impact on economy, land management and land use in those regions… or, you’d hope it would, anyway (who knows, given the current state of environmental protection in the USA). The hypothetical discovery of such an animal would also be regarded by many as one of the biggest ‘wins’ ever scored against ‘establishment science’, and thus could well be a bad thing (viz, “if scientists were wrong about this, what else could they be wrong about?”). And I’ll stop there before we dive into a rabbit-hole of conspiracy theories and coverups.

A big, flightless passerine. The majority of living bird species – over 60% of them – are passerines, or perching birds. This is the great group that includes crows, thrushes, warblers, finches, sparrows and so many others. For all their success, wide distribution and diversity, passerines are generally quite samey. There are no big, long-legged wading passerines, or heavy-bodied diving passerines or flightless running passerines, for example. Why this is so remains mysterious: passerines didn’t take to those niches because… well, they just didn’t. Does this mean that they couldn’t? As usual, we can come up with a few reasons as to why they were ‘constrained’ in evolutionary potential, but any one of those reasons could be overturned by some evolutionary deviant that refuses to pay attention to the rules.

Passerine birds are diverse, to a degree… here’s just a sample of their diversity. This is part of a giant montage that’s being built for   my in-prep textbook The Vertebrate Fossil Record  . Image: Darren Naish.

Passerine birds are diverse, to a degree… here’s just a sample of their diversity. This is part of a giant montage that’s being built for my in-prep textbook The Vertebrate Fossil Record. Image: Darren Naish.

And thus I submit that a particularly large, wholly flightless, cursorial passerine should make itself known to the world. It should be a record-holder as goes size, but not necessarily be that much bigger than the largest known passerines (like lyrebirds and ravens): I’m talking about a bird that weighs 3-5 kg and is thus similar in size to a large chicken. It should be a big, long-legged rail-babbler, quail-thrush or similar, and hence be a denizen of Wallacea or nearby.

Eupetes , the Malaysian rail-babbler. A hypothetical big, flightless passerine should be a close relative of this bird. Image: Francesco Verronesi, CC BY-SA 2.0 ( original here ).

Eupetes, the Malaysian rail-babbler. A hypothetical big, flightless passerine should be a close relative of this bird. Image: Francesco Verronesi, CC BY-SA 2.0 (original here).

A few recently extinct, island-dwelling passerines were flightless, so we do know that passerines have the evolutionary potential to follow this pathway. Such species (a bunting and a few New Zealand wrens… and possibly a few others) were all small (less than 40 g).

A western Asian giant salamander. Giant salamanders (cryptobranchids) are restricted today to eastern Asia (where Andrias occurs) and North America (where Cryptobranchus occurs). Hunting, human disturbance, habitat loss and deterioration, climate change and other issues are putting them into perilous decline, right at the same time as we’re discovering that some of them are species complexes. They were more widespread in the past than they are today, since fossils show that Andrias salamanders were widespread across Europe and Asia between about 28 and 2 million years ago.

An Asian giant salamander ( Andrias ) photographed in captivity. Record-holding specimens of  Andrias  can be 1.8 m long and exceed 60 kg, and some extinct species reached even larger sizes. Image: Markus Bühler.

An Asian giant salamander (Andrias) photographed in captivity. Record-holding specimens of Andrias can be 1.8 m long and exceed 60 kg, and some extinct species reached even larger sizes. Image: Markus Bühler.

While there are very good reasons for the decline and extinction of the animals in the areas concerned, some of the regions where they formerly occurred still have what look like suitable habitat today and are sparsely populated by people. Furthermore, extinct giant salamanders weren’t all denizens of fast-flowing, highly oxygenated streams like those inhabited by the modern populations. Some inhabited ponds and lakes. Ergo: I would really, really like there to be a west Asian cryptobranchid that comes from a habitat considered weird for the other living members of the group. And it doesn’t have to be a giant of 2 metres or more. A hellbender-sized species of 70 cm or so will do fine thank you very much.

Some extinct cryptobranchids - this is Zdeněk Burian’s reconstruction of  Andrias scheuchzeri  - inhabited European ponds and lakes. I’ve previously criticised this image for showing the animal as terrestrial. Since then, the proposal has been made that some extinct cryptobranchids (albeit not  A. scheuchzeri ) were significantly more terrestrial than living species. Image: (c) Zdeněk Burian.

Some extinct cryptobranchids - this is Zdeněk Burian’s reconstruction of Andrias scheuchzeri - inhabited European ponds and lakes. I’ve previously criticised this image for showing the animal as terrestrial. Since then, the proposal has been made that some extinct cryptobranchids (albeit not A. scheuchzeri) were significantly more terrestrial than living species. Image: (c) Zdeněk Burian.

Again, this is an area of special interest to cryptozoologists, since there have been occasional suggestions that stories, engravings and such from western Asia might reflect folk knowledge of unusually big salamanders in the region. In reality, the images and stories concerned are super-ambiguous and more likely refer to otters and god knows what else.

At left:  Andrias  skull. Image: Darren Naish. At right: Japanese giant salamander ( A. japonicus ) illustration by Y. de Hoev from 1887. Image: Y. de Hoev, public domain ( original here ).

At left: Andrias skull. Image: Darren Naish. At right: Japanese giant salamander (A. japonicus) illustration by Y. de Hoev from 1887. Image: Y. de Hoev, public domain (original here).

A living albanerpetontid. Everyone knows that there are three main groups of living amphibian: caecilians, salamanders and anurans (frogs and toads). But until (geologically) recently, there was a fourth group: the albanerpetontids, sometimes termed albies by those who work on them. Albanerpetontids were geographically widespread, their range including Eurasia, northern Africa and North America, and they were geologically long-lived too. The oldest are from the Middle Jurassic while the youngest are… well, we’ve known of Miocene fossil albanerpetontids for decades, have known of Pliocene specimens since 2005 (Venczel & Gardner 2005), and now know that at least one species persisted into the Pleistocene (Villa et al. 2018). The fact that their fossil record has been creeping towards the Recent means that the possibility of fossil and even extant Holocene specimens being discovered isn’t ridiculous, especially given the small size of these animals and hence tiny size of their bones.

New salamander species are occasionally discovered in Europe and Asia even now. It would be amazing if an animal suspected to be a ‘new salamander’ one day turned out to be a living albanerpetontid. These reconstructions were published by McGowan & Evans (1995). They might have erred in implying that the scales would be externally visible as shown here; more likely is that they were concealed by epidermis, as in other scaly fossil amphibians. Image: McGowan & Evans (1995).

New salamander species are occasionally discovered in Europe and Asia even now. It would be amazing if an animal suspected to be a ‘new salamander’ one day turned out to be a living albanerpetontid. These reconstructions were published by McGowan & Evans (1995). They might have erred in implying that the scales would be externally visible as shown here; more likely is that they were concealed by epidermis, as in other scaly fossil amphibians. Image: McGowan & Evans (1995).

To be frank, a live albanerpetontid wouldn’t be a particularly spectacular animal: it would be a tiny, slim, salamander-like amphibian less than 10 cm long, and it wouldn’t be much fun to watch since it would spend most of its time hiding and burrowing in leaf litter. But among herp-nerds it would be a huge deal. Live albanerpetontids were scaly-skinned (though the scales were not necessarily visible externally), with eyelids, and with adaptations in the snout, skull-roof, neck and body shape linked to head-first burrowing (McGowan & Evans 1995).

An artistic reconstruction of a live albanerpetontid…   produced for my in-prep The Vertebrate Fossil Record  . Image: Darren Naish.

An artistic reconstruction of a live albanerpetontid… produced for my in-prep The Vertebrate Fossil Record. Image: Darren Naish.

A Eurasian palaeognath. Palaeognaths are the big, flightless ratites (ostriches, emus and so on), the superficially gamebird-like, flight-capable tinamous, and their extinct relatives. A huge amount has been written about the evolutionary history and biogeography of these birds, since their distribution is curious and has resulted in all kinds of different models about how they might have spread around the world. I’ve written about this issue at length on previous occasion (the articles concerned being famous for generating the longest-ever comment threads in the history of TetZoo… though all of this is mostly wasted now, what with SciAm’s paywalling of the site, sigh). Living palaeognaths are absent from Eurasia, despite the former present in the region of ancient, flight-capable Paleogene taxa, extinct ostriches and others.

As this map shows, modern palaeognaths occurred everywhere until recently (except Antarctica) with the exception of northern North America and the cooler parts of Eurasia. Extinctions across Eurasia, Madagascar and New Zealand of course saw the disappearance of various members of the group. Image: Darren Naish.

As this map shows, modern palaeognaths occurred everywhere until recently (except Antarctica) with the exception of northern North America and the cooler parts of Eurasia. Extinctions across Eurasia, Madagascar and New Zealand of course saw the disappearance of various members of the group. Image: Darren Naish.

The fact that Paleogene Europe was home to many bird groups that no longer occur there but are now denizens of tropical regions elsewhere leads me to hope for a living palaeognath – a tinamou- or bustard-sized species – that descends directly from archaic Paleogene taxa and now lives in the Asian tropics. It should be a cryptic generalist with barred plumage and a mid-length bill and a reduced flight ability.

A gigantic, predatory, limbed amphisbaenian. Regular readers of TetZoo might know that I really like amphisbaenians: the mostly limbless, bullet-headed ‘worm lizards’ of the American tropics, Africa, and parts of southern Europe and western Asia. Amphisbaenian evolutionary history and biogeography has become increasingly complex in recent years as we’ve learnt a bunch of new stuff about their fossil history, genetics and anatomy. Among the weirdest of amphisbaenians are the ajolotes (or bipedids), the only extant group to possess limbs. These limbs are not small stumps or flaps (as they are in some other near-limbless, serpentine squamates) but well-developed, clawed forelimbs. According to some phylogenetic models, ajolotes are not the sister-group to limbless amphisbaenians but deeply nested within the limbless clade (Conrad 2008, Videl et al. 2008), in which case their limbedness – if you will – perhaps evolved from limbless ancestors. Add to this the fact that some amphisbaenians are robust-jawed, short-faced predators of vertebrates that ambush prey from beneath the surface and bite chunks from the bodies of surface-dwelling mammals and reptiles.

Bipes , an ajolote of Mexico (they might occur in parts of the USA as well). Three extant species are recognised. Image: Darren Naish.

Bipes, an ajolote of Mexico (they might occur in parts of the USA as well). Three extant species are recognised. Image: Darren Naish.

So then… where oh where are the giant, limbed, robust-skulled, vertebrate-eating amphisbaenians? By ‘giant’, I am not talking about a graboid-sized monster of several metres (though that would be nice), but a more reasonable animal of a mere 1.5 metres or so. Easily the stuff of nightmares. They could inhabit warm regions of any continent.

Carl Gans’s illustration of a burrowing ajolote, showing how the large, well-clawed forelimbs function in propulsion. This is clearly a Five-toed worm lizard  Bipes biporus ; the other extant species have four and three digits, respectively. Image:   Gans (1974)  .

Carl Gans’s illustration of a burrowing ajolote, showing how the large, well-clawed forelimbs function in propulsion. This is clearly a Five-toed worm lizard Bipes biporus; the other extant species have four and three digits, respectively. Image: Gans (1974).

Is there any reason to think that gigantic, predatory, limbed amphisbaenians might actually exist and await discovery? Nope. But I wish it were so. Regular readers might recognise that such creatures are denizens of the alternative-timeline Earth of the Squamozoic, but I’m sure that that’s coincidental.

What would a gigantic, predatory, limbed amphisbaenian look like? Like this, of course. Image: Darren Naish.

What would a gigantic, predatory, limbed amphisbaenian look like? Like this, of course. Image: Darren Naish.

An African or west Eurasian, long-beaked river dolphin. On several occasions within the history of odontocete cetaceans (‘toothed whales’), lineages have moved into brackish and estuarine environments, and eventually made the transition to committed freshwater life. There are the Asian Platanista species, the recently extinct Lipotes of China, and the tropical American Inia species. Once united within Platanistoidea and thought to be close kin, we know today that these animals represent at least three separate transitions to the freshwater environment (the term Platanistoidea is now restricted to the Platanista lineage alone). In addition, members of other groups – I’m thinking of the delphinid Orcaella – occur in rivers within parts of their range. There’s also a fossil beaked whale that might be indicative of freshwater specialisation in yet another odontocete group (Mead 1975).

River dolphins are pretty special looking. This is a Ganges river dolphin ( Platanista gangetica ). Image: Zahangir Alom / Marine Mammal Commission / National Oceanic and Atmospheric Administration, public domain ( original here ).

River dolphins are pretty special looking. This is a Ganges river dolphin (Platanista gangetica). Image: Zahangir Alom / Marine Mammal Commission / National Oceanic and Atmospheric Administration, public domain (original here).

In view of all this, why aren’t there river-dwelling dolphins in Africa, Europe or western Asia? Again, the answer seems to be… there just aren’t. A few fossil taxa suggest that such animals might have evolved if things had gone another way (there are fossil platanistoids from the Caucasus, for example). But I humbly submit that the great river systems of tropical Africa, the Tigris-Euphrates system of western Asia and the Danube, Po, Ebro, Dniester and others of southern Europe would be much improved if only we knew of their endemic riverine dolphins. I’m talking about a true riverine specialist, convergent with Inia and Platanista, with a long beak, spike-like teeth, reduced eyesight, the works. And if you want to play fast and loose with antiquarian literature and anecdote, there are references in the literature to ‘river dolphins’ in the Nile and there are even one or two eyewitness accounts from central Europe that describe long-beaked ‘dolphins’ seen in rivers and lakes.

If there are extant west Eurasian or African river dolphins, they should look like this. This is a hypothetical species, modelled on the American  Inia  and Asian  Platanista . Image: Darren Naish.

If there are extant west Eurasian or African river dolphins, they should look like this. This is a hypothetical species, modelled on the American Inia and Asian Platanista. Image: Darren Naish.

An endoparasitic tetrapod. Tetrapods have become parasites on several occasions. Vampire bats are parasites of birds and mammals, and it’s even been argued that some blood- and milk-eating human populations can be considered parasites of the mammals they rely on (though the mammals concerned are domesticated, so it’s complicated). Elsewhere among vertebrates, everyone knows about the parasitic catfishes that invade the gills of other actinopterygian fishes and even the urethras of mammals; less familiar is the fact that other actinopterygians can, on rare occasion, become trapped inside the bodies of other vertebrates and then make a successful living. Yes, you read that right. I have in mind the case where two Snubnosed or Pugnose eels Simenchelys parasitica were discovered living inside the heart of a mako shark (Caira et al. 1997; see also Eagderi et al. 2016). This eel is not – despite its name* – ordinarily an internal parasite: this was a case of facultative endoparasitism!

At left: a snubnosed eel found living inside the heart of a shark. Eels are not tetrapods, it’s true. But here’s evidence that aquatic vertebrates can become endoparasites. Image: Caira  et al . (1997). At right: an aquatic typhlonectid caecilian. Surely it’s only a matter of time before we discover an endoparasitic one of those as well. Image: Neil Phillips.

At left: a snubnosed eel found living inside the heart of a shark. Eels are not tetrapods, it’s true. But here’s evidence that aquatic vertebrates can become endoparasites. Image: Caira et al. (1997). At right: an aquatic typhlonectid caecilian. Surely it’s only a matter of time before we discover an endoparasitic one of those as well. Image: Neil Phillips.

There are all kinds of reasons why a tetrapod couldn’t become an endoparasite, respiration being high on the list. A hypothetical endoparasitic tetrapod would have to be small, with remarkable tolerance of unusual chemical and thermal conditions, with low oxygen requirements, and most likely with the ability to respire cutaneously or via gills. In other words, it should be the world’s weirdest caecilian. As if caecilians aren’t weird enough, I’d love there to be small, endoparasitic caecilians. Given that some caecilians are already aquatic gill-breathers that will consume the tissues of fish (exhibit A: the sequence from River Monsters where Jeremy Wade discovers swarming typhlonectid caecilians in the carcass of a large fish), I predict these animals to be aquatic, South American species that parasitise actinopterygians and aquatic mammals, like Inia the river dolphin.

* Snubnosed eels were given the name ‘parasitica’ because they opportunistically latch on to the bodies of larger fish and eat away at the flesh. They were not thought to ever be proper internal parasites prior to 1992.

And that’s where I’ll stop for now. I actually came up with a list containing numerous additional ‘wish list’ animals but time is against me. Maybe I’ll cover them in another article. Whatever, this was all a bit of fun and I hope you enjoyed it.

For TetZoo articles relevant to the issues covered here, see…

PS I’m going to stop linking to the SciAm run of TetZoo articles soon, because I cannot access them at all and they’re now all but useless. They all need to be relocated to an open-access site.

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.

Refs - -

Caira, J. N., Benz, G. W., Borucinska, J. & Kohler, N. E. 1997. Pugnose eels, Simenchelys parasiticus (Synaphobranchidae) from the heart of a shortfin mako, Isurus oxyrinchus (Lamnidae). Environmental Biology of Fishes 49, 139-144.

Conrad, J. 2008. Phylogeny and systematics of Squamata (Reptilia) based on morphology. Bulletin of the American Museum of Natural History 310, 1-182.

Eagderi, S., Christiaens, J., Boone, M., Jacobs, P. & Adriaens, D. 2016 Functional morphology of the feeding apparatus in Simenchelys parasitica (Simenchelyinae: Synaphobranchidae), an alleged parasitic eel. Copeia 104, 421-439.

Gans, C. 1974. Biomechanics: An Approach to Vertebrate Biology. J. B. Lippincott Company, Philadelphia, Toronto.

Graham, G. L. 2002. Bats of the World. St. Martin’s Press, New York.

Mackal, R. P. 1987. A Living Dinosaur? In Search of Mokele-Mbembe. E. J. Brill, Leiden.

McGowan, G. J. & Evans, S. E. 1995. Albanerpetontid amphibians from the Cretaceous of Spain. Nature 373, 143-145.

Mead, J. G. 1975. A fossil beaked whale (Cetacea: Ziphiidae) from the Miocene of Kenya. Journal of Paleontology 49, 745-751.

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

Simons, E. L. & Ettel, P. C. 1970. GigantopithecusScientific American 222 (1), 77-84.

Venczel, M. & Gardner, J. D. 2005. The geologically youngest albanerpetontid amphibian, from the Lower Pliocene of Hungary. Palaeontology 48, 1273-1300.

Vidal, N., Azvolinsky, A., Cruaud, C. & Hedges, S. B. 2008. Origin of tropical American burrowing reptiles by transatlantic rafting. Biology Letters 4, 115-118.

Villa, A., Blain, H.-A. & Delfino, M. 2018. The Early Pleistocene herpetofauna of Rivoli Veronese (Northern Italy) as evidence for humid and forested glacial phases in the Gelasian of Southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology 490, 393-403.

The Dougal Dixon After Man Event of September 2018

“Speculative biology, or speculative evolution, is a term that refers to a very hypothetical field of science that makes predictions and hypotheses on the evolution of life in a wide variety of scenarios and is also a form of fiction to an extent. It uses scientific principles and laws and applies them to a "what if" question” — the Speculative Evolution Wiki

Dixon 1981; Dixon 2018.

Dixon 1981; Dixon 2018.

On September 11th 2018, I had the extraordinary privilege of appearing on stage with author, artist, editor, model-maker and visionary Dougal Dixon to discuss his famous book of 1981 After Man: A Zoology of the Future (Dixon 1981). We were joined by more than 215 interested members of the public, effectively filling the venue (Conway Hall in London).

At Conway Hall once again, such a noble venue. Image: Will Naish.

At Conway Hall once again, such a noble venue. Image: Will Naish.

As you might have gathered if you’re a regular Tet Zoo reader, After Man has just been republished, and our on-stage event – hosted by New Lands London, and arranged by Scott Wood – also served as the launch for the new, 2018 edition (Dixon 2018). This was on sale and available for signing at the meeting.

As you can see from these photos, Dougal brought along a treasure trove of material relevant to the genesis of After Man, including his original sketches, text, draft spreads and the original page plan for the entire work. The latter consisted of large card mock-ups with small, rectangular vignettes depicting the planned look for each page. Dougal explained how he took these documents along to two – yes, just two – publishers and immediately got the green light from both. Evidently, he knew exactly what he was doing.

Dougal on stage, showing the original page plan for  After Man . Small vignettes, showing the planned look of all the pages, are arranged in sequence. Image: Darren Naish.

Dougal on stage, showing the original page plan for After Man. Small vignettes, showing the planned look of all the pages, are arranged in sequence. Image: Darren Naish.

The original sketches are excellent and a testament to Dougal’s skill and planning. The quality of these illustrations also leave you wondering why it isn’t Dougal’s art that we see in the final book, and I impressed upon him during our discussion how fantastic it would be to one day see these unpublished illustrations in another book: a ‘The Making of After Man’ or something along those lines. We’ll see.

Foreign language translations of  After Man , a Vortex and Raboon model, and relevant magazine issues (like the October 1981 ish of  BBC Wildlife ). (c) Dougal Dixon. Image: Darren Naish.

Foreign language translations of After Man, a Vortex and Raboon model, and relevant magazine issues (like the October 1981 ish of BBC Wildlife). (c) Dougal Dixon. Image: Darren Naish.

Several of the creatures pictured on the draft spreads were obvious prototypes of versions that made it into the final book but others were evidently abandoned at some point. A few of those ‘prototypes’ showed how the original animals had a different look relative to the published descendants: the bone-cracking Ghole Pallidogale nudicollum, to take one example, looked a lot more like a big mongoose in its original guise than is obvious in the book.

Original text and original draft double-page spread for  After Man , showing creatures inhabiting tropical grasslands. You’ll recognise some (but not all!) of the creatures as the prototypes of versions that made it to final publication. (c) Dougal Dixon. Image: Darren Naish.

Original text and original draft double-page spread for After Man, showing creatures inhabiting tropical grasslands. You’ll recognise some (but not all!) of the creatures as the prototypes of versions that made it to final publication. (c) Dougal Dixon. Image: Darren Naish.

Dougal also brought foreign-language editions of After Man and a few models of After Man’s creatures with him, including a resin model of the Vortex Balenornis vivipara (a reasonable number were made, but it may be that this is the only one still in existence) and a wonderfully detailed Desert leaper Aquator adepsicautus. Alas, the Night stalker Manambulus perhorridus model pictured on the dustjacket of Dixon 1981 – the model I most wanted to see – is not in Dougal’s possession so was a no-show.

Desert leaper model. For the handful of you that haven’t read  After Man , the Desert leaper is a giant, desert-dwelling muroid rodent (in cases more than 3 m long) that undergoes significant fluctuation in fat deposition (and hence mass) according to season. (c) Dougal Dixon. Image: Darren Naish.

Desert leaper model. For the handful of you that haven’t read After Man, the Desert leaper is a giant, desert-dwelling muroid rodent (in cases more than 3 m long) that undergoes significant fluctuation in fat deposition (and hence mass) according to season. (c) Dougal Dixon. Image: Darren Naish.

As goes our actual discussion, we covered the backstory to After Man (some of which will be familiar if you know the interview I published at ver 3 back in 2014), the response from critics and reviewers and the many overseas trips Dougal got to enjoy as a consequence of the book’s success, the substantial interest from Japanese markets and the Japanese stop-motion and animated movies (we watched a short segment from the stop-motion movie, copies of which were given away on DVD to people buying the book), the various efforts by studios in Hollywood to get an After Man movie off the ground, and the connection between After Man and Dougal’s more recent project Green World (thus far only published in Japanese).

Original sketches, by Dougal, of creatures illustrated for  After Man . The animals were then re-illustrated by various other artists. (c) Dougal Dixon. Image: Darren Naish.

Original sketches, by Dougal, of creatures illustrated for After Man. The animals were then re-illustrated by various other artists. (c) Dougal Dixon. Image: Darren Naish.

We finished with a Q&A session and audience participation. Questions included the ‘new look’ Night stalker (yup… I shall say no more), Dougal’s thoughts on the future of humanity, how and which fossil species had influenced the creatures of After Man, and what might be different in After Man if Dougal were to write the book today.

The Vortex model that Dougal brought along. It’s about 60 cm long. (c) Dougal Dixon. Image: Darren Naish.

The Vortex model that Dougal brought along. It’s about 60 cm long. (c) Dougal Dixon. Image: Darren Naish.

For a lifelong fan of After Man and Dougal’s connected writings, this event was an absolute thrill and I’m tremendously happy to have been involved. And judging by our audience’s response, it was enjoyed by everyone who attended too: thanks so much to everyone who came along and participated.

Two final things are worth saying. Firstly, I was asked innumerable times whether the event was going to be recorded. Alas, I was simply unable to organise this or even remember it given all the other stuff I had to worry about, though I think (and hope) than an audio recording exists. Secondly, this is not the only Dougal Dixon-themed event of 2018! Dougal is also on stage at this year’s TetZooCon when he will be joined by Gert van Dijk (of Furahan Biology and Allied Matters) in a discussion on speculative biology. TetZooCon happens on Oct 6th and 7th in London and tickets are still available. Dougal will also be bringing archive material to that meeting as well!

Darren Naish (l) and Dougal Dixon (r) on stage at Conway Hall, September 2018. Image: Will Naish.

Darren Naish (l) and Dougal Dixon (r) on stage at Conway Hall, September 2018. Image: Will Naish.

My thanks to Dougal for being such a brilliant person to talk to and for all the material he brought along, to Scott and everyone else at Conway Hall and New Lands for organising things and setting it all up, to the Breakdown Press people for the book selling, and to our brilliant audience for their interest, enthusiasm and participation.

For previous articles on speculative biology, see…

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

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Dixon, D. 1981. After Man: A Zoology of the Future. Granada, London.

Dixon, D. 2018. After Man: A Zoology of the Future. Breakdown Press, London.