Notwithstanding the occasional penguin or spinosaur, Dinosauria is a clade predominantly composed of land-living animals, adapted towards terrestrial habitats and diets (and if you are a reader of this blog you are probably smart enough to know that plesiosaurs, mosasaurs and such were not dinosaurs). There are numerous pieces of evidence, anatomical, palaeobiological, coprolitic and especially stratigraphical that attest this. However, it took some time to come to this consensus and if you are a little older you may very well remember old books depicting a few select groups as amphibious. This mainly concerned the sauropods and the hadrosaurs, which used to be imagined as living in swamps and feeding on soft aquatic plants, due to supposed deficiencies observed in their anatomy. These interpretations largely vanished with the Dinosaur Renaissance of the 70s and 80s, which reinterpreted dinosaurs as highly competent animals on land that occupied the same ecological niches as modern mammals would.
![]() |
Fig. 1: Corythosaurus, if I had drawn it according to one Martin Wilfarth... which I did. |
Nonetheless, the idea of aquatic dinosaurs did have somewhat of a comeback in the 2010s, due to writer Brian J. Ford and his book Too Big to Walk. As the title implies, Ford, who is not a paleontologist and self-admitted that he is naïve about dinosaurs, thinks that all of them were not actually capable of living and reproducing on land and instead lived their whole lives wading through water on a rather alien prehistoric Earth dominated by balmy, shallow seas with very little dry land. But this revival, despite initial media attention, was thankfully more of a stillbirth. Ford’s arguments mainly stemmed from incredulity and ignorance on modern dinosaur research, seemingly with zero awareness that aquatic dinosaurs have already been considered and disproven by multiple generations of scientists before him, his book is poorly written and spends most of its length not even discussing dinosaurs (Naish 2024). Instead, it is overly occupied with self-aggrandization of the author about how revolutionaries like himself are rarely ever taken seriously in their times. To paraphrase Carl Sagan, people may have laughed at Galileo, but they also laughed at Bozo the Clown. There is very little worthwhile content in Ford’s body of work and it is widely seen as a prime example sensationalized pseudoscience.
The one positive of this stupidity I have been able to find though is that the authors critiquing Ford have brought back into attention one Martin Wilfarth, a German researcher mainly publishing during the 1930s and 50s, who, as some of them (Naish 2024) have noted, already prefigured aspects of Ford’s thesis over half a century ago (without Ford ever acknowledging him), as he was the first to propose that all dinosaurs had aquatic adaptations in some form. But a bit of a shocking thing occurred to me recently, when, out of curiosity, I actually read an original German copy of Wilfarth’s 1949 book Die Lebensweise der Dinosaurier (“The Lifestyle of the Dinosaurs”). Wilfarth was decidedly not a proto-Brian-Ford. No, he was an anti-Brian-Ford. Though he ultimately came to the wrong conclusions, based on wrong assumptions about the Mesozoic environment, Wilfarth’s writings do demonstrate a remarkable observational skill and intuition about dinosaur anatomy, as well as competent knowledge of the state of dinosaur research during his time. While he got some things wrong, he also got quite a few notable things very right, often decades ahead of other paleontologists.
Despite that, almost none of the authors I could find discussing Wilfarth meaningfully engage with this fact. Instead, they either present only very brief summaries of his theses or outright insult the man’s work, often with confusing errors in their reading of him. Edwin Colbert’s (1950) review of Wilfarth’s book is less than two paragraphs long and basically ends with him calling it garbage without elaboration. Charles Sternberg (1939) criticized Wilfarth on the basis of thinking that he proposed that Parasaurolophus had a food-gathering, elephant-like trunk, which, as we will see, Wilfarth never actually did. Adrian Desmond (1975, p. 52) repeats this false claim and seems to have only read said response by Sternberg. The same seems to be true for Knoll et al. 2004, who also seem to cite Wilfarth second-hand through Sternberg. Darren Naish (2024) commendably went through the effort of actually including illustrations from Wilfarth (1949a) in his paper discussing Ford, though in the image description he expresses surprise as to why Wilfarth would depict Iguanodon and Kentrosaurus as aquatic. Unless Naish is not one to indulge in hermeneutics, I do not see why those images would be surprising, given the fact that Wilfarth extensively argues in the text why he thinks these animals were aquatic.
All of these authors are English-speakers and I do not know whether they had or have any German proficiency. Wilfarth’s works have likewise never been translated into English. It may be a grave accusation that I may not be able to prove, but I have a sneaking suspicion that perhaps none of these people have actually directly read the texts written by Wilfarth and have only engaged with him through second-hand commentaries, summaries, guesswork or illustrations. In the process, a lot of his work, I think, has ended up overly simplified, if not outright misrepresented. For example, Benton 1990 lists Wilfarth as assuming dinosaurs were underwater organisms, while Naish 2024 basically equates Wilfarth’s ideas with those of Ford. This is only correct in the broadest of terms and I think gives a false impression. Unlike Ford, Wilfarth never claimed that any dinosaurs were fully marine or were too heavy to have been able to walk on land. He specifically interpreted all of them as specialists of the tidal flats, neither fully marine nor fully terrestrial, but perfectly amphibious (or “amphibiotisch”, as he says in that wonderful old Science German), meaning they needed to be able to walk during ebb and egg-laying, with some groups being more terrestrial than others.
The purpose of this article here is twofold. Mainly, I want to present a translated overview of Wilfarth’s complete vision of the Mesozoic. While this vision is ultimately wrong and I do not wish to rehabilitate it as somehow viable, I still want to relativize it and give Wilfarth a fair evaluation. Even if his conclusions ended up flawed, the data and points he raised in the process of coming to those conclusions were still remarkably observant, often highlighting features of dinosaur anatomy that many others at the time ignored. While at first glance his idea of aquatic dinosaurs seems regressive, Wilfarth proves quite unique for a paleontologist from the 1940s for actually being very charitable to our favorite megafaunal reptiles, viewing them not as evolutionary failures as many others did, but actually as quite sophisticated and ingeniously adapted for their environment and ecology, ultimately going extinct due to no fault of their own. All too often in the history of science, people are reduced down to their greatest mistakes, without appreciating their successes. Swiss naturalist Johann Jakob Scheuchzer was one of the founders of crystallography and also perhaps the first person in Early Modern Europe to show that fossils were remains of lifeforms, not lusus naturae, making him arguably the first modern paleontologist. But chances are you only know him as that funny guy who thought a salamander-fossil from Öhningen was a human that died in Noah’s Flood.
The second purpose is the pure fun of hermeneutics, meaning here seeing things from another person’s point of view. Wilfarth’s vision of the Mesozoic is extensive, covering all the then-known dinosaur groups, and quite detailed in his descriptions of speculative life habits and ecosystems. And while it is ultimately based on false assumptions, there is a clear and understandable method to the madness, an internal logic that is at times undeniably clever. While this version of the Mesozoic never existed on our Earth, who is to say that something like it may not exist on an alien planet or on an alternative timeline of Earth? What I am trying to say here is that, if nothing else, Wilfarth will prove a treasure trove of inspiration for worldbuilders and speculative evolution enthusiasts. I may very well incorporate some of his ideas into my own spec works.
As a sidenote, I believe I am the first person online to talk about Wilfarth to this extent and also the first to upload some scans of his fascinating illustrations. So, if in the coming months you see any particular youtubers suddenly talking about Wilfarth, you will know where they got their info from. If you are one of those people, I will know. I have friends everywhere.
Due to the sheer amount, I will have to cut this post up into multiple parts. I have already written all of them and will post them over the span of a few days, so no worries. If you wish to go ahead and read the book yourself, I took the liberty to scan it and thus provide it here for you through this link.
Who is Dr. Martin Wilfarth?
This has been a surprisingly difficult question to answer, because outside of his own papers/books and the few superficial commentaries on those, there is almost no biographical information on him available anywhere, let alone photographs. Almost any online article that mentions him basically describes him as just “that weird German guy who thought Parasaurolophus had a trunk”. Thankfully the cover page of his 1949 book includes the important piece of information that he was born on the 5th December of 1893, in Bernburg, a city in what was, at the time, in the Free State of Anhalt, which in 1947 became part of Sachsen-Anhalt. An address book from 1941 does indeed list a certain Dr. Martin Wilfarth still living in that city. I wonder if he is a relative of the more famous Hermann Wilfarth, a chemist also born in Bernburg, who was involved in the discovery of how plants draw nitrogen from the air. What exactly our Wilfarth was a doctor in, and with what institution he was affiliated, I have not been able to find out for certain, though based on his oldest publications in the 30s, I believe he was a geologist by trade. These oldest papers all concerned odd geological anomalies in the Permian and Triassic rocks of Germany, which he sought to explain with his Great Tides model. In 1938 he released his first paper on dinosaurs, specifically sauropods, and how they fit into the Great Tides model. What then followed was a quite regular stream of publications on hadrosaur facial anatomy, again in relation to tidal life. Wilfarth was able to publish his papers in official journals of German geological societies, and his monographs were produced by respectable publishers like Schweizerbart, indicating that he was not treated as a crank by his peers and did meet the scientific standards of his time and place. The university library of Freiburg im Breisgau contains at least two letters sent by Wilfarth to famous paleontologist Friedrich von Huene, one before, one after the war. I do not know what the content of those letters is, as they were not digitalized, but if one of you readers is from Freiburg, maybe check them out and tell me. At the very least it is interesting to know that these two were in contact with each other and perhaps Wilfarth derived some or most of his dinosaur knowledge from Huene.
Fig. 3: An early hadrosaur with an extendable breathing trunk, somewhat similar to those seen in softshell turtles. More on Wilfarth's hadrosaurs in part 3 (Source: Wilfarth 1949a, p. 75). |
After 1940 there is then a hiatus in his publications, for quite obvious reasons. In the introduction to his book, Wilfarth shares with us one of the few autobiographical details in relation to his work, as well as the state of science in post-war Germany:
“For 20 years I have worked on my hypothesis of the greater tides. What I have been able to publish of it so far are merely fragments. Manuscripts are stacking into piles, but the printing has so far been impossible. Before the war, the summary presentation was not yet ripe, during the war printing was not feasible, now again many years will pass before the scientific life can recover again. This is why I want to place with this another fragment of my hypothesis in front of public critique.” (Wilfarth 1949, p. 1, translated by me).
While brief, the impression I get from Wilfarth here is of a man who, while obsessed with his unorthodox hypothesis, was also passionate about science and dinosaurs, trying his best to study them even through Europe’s most destructive catastrophe and wanting to keep the scientific community alive right when it was possible again during reconstruction. In 1947 he was already publishing again on hadrosaurs, which may be a candidate for the first paper on dinosaurs in post-war Germany. Unlike cranks of today, it is also notable that throughout the introduction, Wilfarth (1949, p. 1) keeps emphasizing how, even if he seems overly sure about the correctness of his hypothesis, he wants his writings to be critiqued, as he knows that this is what drives the scientific process through correction and discussion.
While this paints a positive picture of him, I should add the caveat that there is no information on what political beliefs or affiliations he may have had during the rise of the Nazis and World War 2. Predictably, papers on dinosaurs and marine sediments contain very little about the author’s opinion on the NSDAP. There is also no indication whether or not he was ever drafted into the Wehrmacht, though if he had been, that probably would have been an important detail to include in discussing how he could not publish his work. The only observation I have been able to make, is that, in the discussion of dinosaur extinction, he heavily criticizes the idea of racial senescence (Wilfarth 1949, p. 87). This is maybe an indicator that he was not a big fan of scientific racism, though that is admittedly a huge stretch.
Wilfarth continued to publish from the late 40s into the 70s, though at less frequency than before. He also began writing articles on his ideas in popular science magazines like Orion and Prisma. Of interest for cryptozoologists is a 1949 article for Prisma, in which Wilfarth weighs in on the possibility of some non-avian dinosaurs having survived into the modern day, particularly reports from Africa collected by none other than Ivan T. Sanderson. Mentioned by name are cyptids such as Chipekwe and the “water-lion” Coje ya menia (this was before Mokele-mbembe was widely popularized and became a household name). He is interestingly critical of Sanderson, arguing that his characterization of Carl Hagenbeck is flawed and that all the reports seem to describe different types of dinosaurs, which he thinks are unlikely to have all survived into the modern day (Wilfarth 1949b, p. 281). If a group of dinosaurs survived, he argues, it would have been the ceratopsians, as they were one of the last groups in the Cretaceous and do fit some of the reports. However, he does note that none of the claimed sightings seem to mention the characteristic neck-shield (Wilfarth 1949b, p. 282). Wilfarth curiously makes no mention here of his tidal interpretation of dinosaurs.
Fig. 4: The lambeosaurine Corythosaurus with a fleshy proboscis covering its headcrest. More on Wilfarth's hadrosaurs in part 3 (Source: Wilfarth 1949a, p. 78). |
His last publication I was able to find was a book from 1971, which curiously seems to have been about the structure of atoms, a departure from his usual topic of the Great Tides. Regrettably, I have not been able to find out when exactly he died (and I assume he died, because he would otherwise now be 132 years old and therefore made the news). Considering the date of his last publication, the obvious assumption would be that it was somewhen during the 70s, probably at a respectable age above 80, though apparently without ever being able to publish a definitive magnum opus on his whole hypothesis.
The Great Tides
So what are the Great Tides that Wilfarth seems to have been so obsessed with and which were the driver behind his interest in dinosaurs? The origin of the hypothesis seems to have been in the “Buntsandstein War”, as he calls it (Wilfarth 1938a, p. 292). This was an academic dispute in the early 20th century over the origin of the Buntsandstein, a lithostratigraphic unit in Germany that spans the Latest Permian to Middle Triassic. In the simplest of terms, the dispute was over whether the unit was formed in a dry desert or in a marine environment, due to evidence speaking for both (Wilfarth 1934, p. 349). In addition, there was some confusion about the odd periodicity in which layers of salt, gypsum and anhydrite seems to follow each other in regular intervals. Wilfarth was fond of Ernst Zimmermann’s 1913 hypothesis of the Grossfluten, in which the author proposed that a barrier separating the Germanic Basin from the Tethys Sea regularly broke down before somehow being rebuilt again, allowing the water to enter and consequently evaporate again. Somewhat similar to what happened to the Black Sea or Mediterranean in the past. But Wilfarth found it unlikely that such a barrier existed, let alone that it would be able to rebuild itself again by the same processes that destroyed it (Wilfarth 1934, p. 360), and so he instead proposed that the sea came into and then left the basin again through the action of the tides. Unimaginably huge tides, capable of inundating and then leaving dry again a strip of land the size of Germany for multiple months at a time. These giant, prehistoric tides were of course caused by the fact the Moon used to be much closer to the Earth in the past than it is today. With this he went on to also explain other Paleozoic and Mesozoic formations, such as the Zechstein and Solnhofen. Some part of this hypothesis Wilfarth seems to have based on work by German astronomer Friedrich Nölke (1935). If you are curious, the idea that the Moon is slowly distancing itself from Earth due to tidal friction, and therefore used to be much closer in the past, dates back all the way to Immanuel Kant in 1754, but was only astronomically proven in 1882 by George Darwin, son of Charles Darwin (Ekman 1993).
Today we know that the Moon moves away from Earth at a rate of about 3.8 centimetres per year. Using simple math, that means that, say, 200 million years ago, it would have been roughly 7600 kilometres closer than today. This may sound like a lot at first, but is actually only a 1.9% difference compared to the modern distance of 384’400 km. For further help I actually emailed planetary scientist Xakarus Alldredge (who has a lovely Youtube channel reviewing the scientific accuracy of scifi planets) and he calculated that, in order to generate the tidal range predicted by Wilfarth, the Moon would have needed to oribt at a mean distance of 70’182 km. There was no period in Earth’s history where the Moon ever orbited that closely, unless you believe the Earth-Moon system is over 8 billion years old. However, we can only make this judgement through the gift of hindsight, as this exact rate would not be known until the Lunar Laser Ranging Experiment of the Apollo Program in the 1970s. So, the distance of the Moon and the Earth during the Mesozoic, and therefore the effect this would have had on the tides, would have still been in the realm of uncertain speculation during Wilfarth’s time. I think it is therefore fair to say that, while unorthodox, his hypothesis was within the range of what would have been thought possible at the time. And while Wilfarth was incorrect about tides being a major driver of dinosaur evolution, there is still some merit in researching how changing tidal forces through deep time affected life on Earth. Chemical analysis of growth lines in rudist shells from the Western Interior Seaway have revealed that in the Late Cretaceous, around 70 million years ago, tidal forces made a day on Earth last only 23.5 hours and thus a year consisted of 372 days (De Winter et al. 2020). Byrne et al. 2020 also used computer model studies to suggest that changing tide levels in the Silurian and Devonian were a major driver in the evolution of tetrapods.
Fig. 6: Corythosaurus again, though this time rearing up underwater and with a longer "trunk". For more on Wilfarth's hadrosaurs, see part 3 (Source: Wilfarth 1949a, p. 79). |
In this context, it is interesting that Wilfarth saw himself as somewhat of a rebel against actualism, or at least too rigid actualism. While he still denies catastrophism and sees the validity of the actualistic principle of using the present as a key to the past, he thinks the actualist school of geology has itself become a new dogma that is just as damaging (Wilfarth 1949a, p. 1 – 2). According to him, while it is reasonable to assume that the phenomena that shaped the past are the same as the ones that work today, it is not always reasonable to assume that those phenomena always operated within the same scale as they do today (Wilfarth 1949a, p. 2). In his analogy, just because glaciers are building up today according to the same principles as they were in the Pleistocene, nobody would deny that the ice ages did not exist or were a phenomenon with no equal in the modern day. While coal is still being produced today in some bogs, it does not happen anymore at the same scale as it did in the Carboniferous. His Great Tides, he says, are the same phenomenon as the modern tides, just at a different scale. His criticism of actualism is interesting from a modern perspective and I wonder what his reaction would have been to someone like Walter Alvarez. One of the greatest challenges to the idea of a rapid dinosaur extinction caused by an asteroid impact faced in the 1980s was not a lack of evidence but the simple fact that it went against the actualism and gradualism paleontologists at the time insisted on, something frequently criticized by Alvarez. It took some considerable amount of time and evidence to convince them that actualism is not the only principle that has shaped the Earth but that there have also been factors unlike anything observed in the modern day. As Wilfarth (1949a, p. 2) succinctly puts it: “If the past was always like today, then we wouldn’t need to study the past.”
![]() |
Fig. 7: The Wattenmeer of Germany's North Sea, the largest intertidal zone of the world. According to Wilfarth an environment like this must have covered most of Earth's continents in the Mesozoic. (Source: Wikimedia). |
The reception to Wilfarth’s hypothesis within Germany is difficult to gauge. At least one person, geological engineer Ernst Fulda (1937a-b, 1938a-b, 1939), seems to have taken it seriously and frequently cited the Great Tides as a potential explanation for certain salt deposits in the German Triassic. Otherwise Wilfarth seems to have been criticized for not adhering to actualism and also for the fact that, if such giant tides had been a regular occurrence, basically leaving no piece of land permanently dry, then surely every land animal would have drowned. His work on dinosaurs was basically driven as one big response to this latter criticism:
“Sure, rabbit and deer, hamster and ant would have been unthinkable during the time of the Great Tides, but were there such organisms during the Mesozoic? To show that the Mesozoic fauna was indeed compatible with the greater tides, I have devised this reconstruction. Regarding water animals like fish, crocodile, turtle, Plesiosaurus, Ichthyosaurus, etc. I did not have to expect great resistance. The dinosaurs, with their true walking-feet, are what gives consideration. Here the amphibiotic character of these saurians had to be demonstrated. It was my task to show that the terrible lizards could not only endure the great floods , but that their whole uniqueness was formed by the interplay of ebb and flow, that they could only be fully understood through the hypothesis of the greater tides.” (Wilfarth 1949a, p. 3, translated by me).
Wilfarths’s writings on the Great Tides are extensive and I have not been able to read all of them, in part because not even my city’s library had most of them (future updates to this section may therefore be in order). Among the parts that remain vague and confusing are how and why exactly Wilfarth thought the tides lasted multiple months, because, as Xakarus noted to me, the tidal period is tied to the Earth’s rotation, not the Moon’s orbit, and since Earth would have rotated faster in the Triassic the tides would have therefore actually lasted shorter than today.
For now it suffices if we list the basic metrics he provides in Die Lebensweise der Dinosaurier. In the Triassic, the average tidal range was a whopping 30 metres tall and ebb and flow lasted each multiple months (Wilfarth 1949a, p. 3 - 4). As the Moon gradually receded from the Earth, so too did the tides become less extreme. By the beginning of the Late Cretaceous, the tidal range was not more than 3 metres and ebb and flow lasted each 60 hours (Wilfarth 1949a, p. 3 – 4). At the end of the Mesozoic and the beginning of the Cenozoic (Tertiary as it is still called here), our modern values began to take shape, with a range of only 1 metre and ebb and flow each lasting 6 hours. Dinosaur anatomy per Wilfarth must be understood of evolving first in these extreme Triassic conditions and then gradually adapting towards the receding tidal zone.
“The Swimbreathers”
Die Schwimmatmer (Pseudosuchia)
As you can see here (and also fig. 10), Wilfarth tried to give each group a German colloquial name in accordance to their imagined lifestyle. I tried my best to translate these into English, but the results will inevitably sound clunkier than the originals. Pretty standard for the time, Wilfarth (1949a, p. 6) imagines the ancestor of the dinosaurs to have evolved from a small, generally crocodile-like animal from the group that was then called Pseudosuchia (the much-maligned term “thecodont” is not used once by Wilfarth, so props to him). Today this is generally (and ironically) considered the line of archosaurs leading up to the true crocodylians, but in the 40s Pseudosuchia was a much vaguer group thought to only include gracile, sometimes bipedal archosaurs, such as Ornithosuchus, Hesperosuchus and Erpetosuchus, which were originally not thought to be directly related to crocodiles. Instead, they were often interpreted to have been the paraphyletic ancestral stock out of which evolved at least the saurischian dinosaurs (see for example Colbert 1964).
Fig. 9: Wilfarth comparing the pelves of a Ceratosaurus, an alligator and Archaeopteryx. (Source: Wilfarth 1949a, p. 10). |
Wilfarth’s speculative ancestral pseudosuchian was an amphibious quadruped that lived at the edges of the giant Triassic tidal basins. Instead of adapting to the “Dauermeer”, the permanent ocean, and therefore evolving flippers like the ancestors of the plesiosaurs and ichthyosaurs did, this ancestral form remained in the tidal zone and therefore still required its legs so it could walk during ebb (Wilfarth 1949a, p. 6 – 7). During the “Great Ebb”, this animal walked on all fours and hunted sea life stranded in the ephemeral pools, ponds and lakes that formed. As well, it dug up aestivating animals waiting in the mud for the return of the sea, such as lungfish, worms, crabs, clams and brachiopods. When the Great Flood came back, it lived as an aquatic predator of fish very much like a modern crocodile. To breathe, it still actively swam to the top of the water, hence why Wilfarth names it the Schwimmatmer. But as time wore on, these animals found it easier and more energetically efficient to stay in shallow water and, instead of spending energy with swimming, they simply reared their head out of the water by rearing up onto their hindlegs, leading to the origin of the dinosaurs.
The view is here then that dinosaurian bipedalism evolved in an aquatic context. This was not an idiosyncrasy of Wilfarth, but a pretty common idea at the time. Crocodylians in water today may also stand up on their hindlegs when they need to take a breath or just keep their head above the water, and their legs are naturally stronger and a bit longer than their arms due to being used to propel themselves. Together with a thick paddle-tail acting as counterbalance, the early crocodile-like archosaurs were thus seen as perfectly pre-adapted towards bipedalism. This view was also endorsed by Romer (1956) and could be found in popular books all the way into the Dino Renaissance (see McLoughlin 1979 – 1981 for example). The sole difference between these authors and Wilfarth is that they imagined bipedalism evolving in freshwater, not in a tidal setting. Only in the 80s did some (Norman 1985) begin to express doubt and reinterpret bipedalism as evolving in a terrestrial setting.
“The Breathjumpers”
Die Atemspringer (Saltopodidae)
Like most people at the time, Wilfarth uses the hips as a major characteristic of the dinosaurs. While similar from the side, he correctly observes that, viewed from the front, there is a stark difference between the hips of a crocodylian and a typical dinosaur. The crocodile, and in turn the ancestral pseudosuchian, has a shallow “Korbbecken” (Basket-pelvis). As the earliest dinosaurs began using their hindlegs for breathing much more, the pelvis adapted by lengthening the ischia and pubis bones in order to support the longer hindlegs and to become levers for the muscles, allowing the animal to easily swing its spine and tail up and down (Wilfarth 1949a, p. 10 – 12). When on dry land, this allowed for true bipedalism, with the new pelvis and long tail allowing for an easy-to-control balance like a seesaw during walking and running (Wilfarth 1949a, p. 11 – 12). These are all views pretty much in line with our modern understanding of dinosaurs using their stiff spines and tails as counterweights. Which makes it ironic that Wilfarth was actually criticized for this during his time. As Wiman (1942, p. 246) wrote: “Wilfarth’s reconstruction of Parasaurolophus becomes a caricature, apart from the breathing proboscis, by him also imagining the bipedal dinosaurs as seesaws who could not lower their head without raising their tail and vice versa.”
When in water, however, the lever-pelvis also allowed for “breath-jumps”. The first group of true dinosaurs that Wilfarth recognises are the Atemspringer, a family he calls Saltopodidae. Included in this group by him are genera such as Scleromochlus, Saltopus, Saltoposuchus, Aetosaurus and Hallopus, which he says are all characterized by a tubercle on the calcaneum which indicates saltational habits. But, as he says, previous researchers were “actualistically naïve” when they thought these animals were jumping in a terrestrial or arboreal context. Instead, they lived in shallow water during flow and when the water level became too high to stand in, they jumped up to the surface (Wilfarth 1949a, p. 7 -8). Wilfarth further supports this aquatic interpretation by pointing out that the famous Chirotherium ichnofossils of Germany, which he attributes to these animals, do not show a central rut that would have been produced by a dragging tail, therefore indicating that the tail was floating in the water (Wilfarth 1949a, p. 11). This funnily enough mirrors similar arguments made after Roland T. Bird found sauropod tracks at Paluxy River, which likewise were lacking the expected tail-rut (Desmond 1975, p. 125). Today we of course know that there was no tail-rut because muscles and tendons were holding the tail high in the air. Another modern irony is of course that, except for Saltopus, none of Wilfarth’s Saltopodidae are seen anymore as being particularly close to dinosaurs. Scleromochlus was a (lagerpetid?) pterosaur-relative, Saltoposuchus and Hallopus were crocodylomorphs and Aetosaurus was, well, an aetosaur. But there were also other researchers at the time that misinterpreted these animals as being small dinosaurs.
Fig. 12: Wilfarth's explanation for the dinosaurian pelvis as being used as an efficient seesaw that allowed poikilotherm animals to achieve bipedal balance (Source: Wilfarth 1949a, p. 11). |
Speaking of classification, Wilfarth identifies the evolution of a “Hebelbecken” (lever-pelvis) as the central distinguishing characteristic of the Dinosauria that makes them different from all other saurians with more primitive basket-pelves. “The name dinosaurs (terror saurians) does not say much, as one can also be terrified by a crocodile. It would be better to call the dinosaurs ‘lever-pelvis-saurians’ [Original: Hebelbeckensaurier], if habit and the law of priority did not stand in the way.” (Wilfarth 1949a, p. 10, translated by me). Most notably, Wilfarth indicates that this lever-pelvis evolved only once and that the Ornithischia evolved from a dinosaur that already had a saurischian lever-pelvis (Wilfarth 1949a, p. 52 – 55). In modern cladistic terms, he views the lever-pelvis as the synapomorphy of a monophyletic Dinosauria, meaning the dinosaurs are a natural group with a single common ancestor. I hold this to be significant, because most leading experts at the time, and even quite after, would have claimed the exact opposite. Ever since Harry Govier Seeley (1887) it had become common to see Saurischia and Ornithischia as two unrelated groups that evolved independently out of different “thecodont” reptiles, making Owen’s Dinosauria an informal term that continued to be used merely out of convenience, not because it was thought to be a natural group. So too argued Friedrich von Huene in 1914. All the way into the 1970s, one could find popular books (McLoughlin 1979 for an example) stating that, technically, there was no such thing as a dinosaur. Wilfarth already favouring dinosaur monophyly in the late 40s is a fact that has so far gone unappreciated.
“The Anchortailers”
Die Verankerungsschwänzler (Podokesauridae)
As the saurischian dinosaurs began exploiting deeper water levels, they evolved longer necks and legs in order to breathe more easily, eventually becoming the Podokesauridae in the Late Triassic, an outdated term for the Coelophysoidea. These Wilfarth imagines as larger, even more specialized versions of the Breathjumpers. Here we really start seeing him be creative with interpreting anatomy. For one, the sclerotic rings he sees as a specific adaptation that allowed the dinosaurs to change their focal length, so that they could see well both on land and in water (Wilfarth 1949a, p. 12). The fact that he even thought of that problem shows that he certainly had an eye for detail.
Fig. 13: Motion study of Podokesaurus, using its curled-up tail as an anchor in the sediment in order to float in water without being carried away by the current. (Source: Wilfarth 1949a, p. 13). |
The name “Anchor-tailers” is given, because Wilfarth (1949a, p. 14) thinks that, when standing or jumping up was not enough anymore to breathe, the thin, long tail-tip of these dinosaurs could curl up and be used as a helical anchor in the sand or mud to allow the dinosaur to passively float in the water “like a balloon” without the danger of being carried away by the tidal current. This was a further adaptation towards changing conditions of the Late Triassic, when tides slowly started to become less extreme, but in turns currents became stronger. Intriguingly, Wilfarth (1949a, p. 8) claims that direct evidence of these anchorings have been found in the form of a trace fossil he calls “Dämonhelix”. I have unfortunately not been able to find out precisely what he is referring to here. Daemonhelix was one of many names for the infamous “devil’s corkscrews” ichnofossils, but these were Oligocene to Miocene burrows from America produced by the prehistoric beaver Palaeocastor. This makes me unsure if Wilfarth was really referring to these, especially also because the illustrated anchor-tail does not line up with the deep corkscrews.
With the concept of the anchor-tail, Wilfarth also states what he believes is another guiding aspect of dinosaur anatomy. He sees all these animals as explicitly poikilotherm (“cold-blooded”) and therefore adapted towards saving as much energy as possible. Throughout the text he repeats the claim that dinosaur evolution always strove to replace active muscle work with passive bone construction (Wilfarth 1949a, p. 14). Also interesting are his notes on dinosaur ontogeny. Laying their eggs on land, the young chicks of dinosaurs would have only gradually adapted to water as they grew (explicitly invoking Haeckel’s discredited idea of ontogeny recapitulating phylogeny) and in the process would have self-stratified, each dino picking a water-level that suited its height as it grew. In some sense this prefigures the modern idea that the young of the largest dinosaurs occupied different ecological niches than the adults as they grew in size. Wilfarth (1938a, p. 290) claims this also explains why juvenile dinosaurs are never found with the adults, referencing Matthew 1915.
“The Small-Fauna-Pluckers”
Die Kleintierpflücker (Plateosauridae)
At this point it is perhaps worth mentioning Wilfarth’s odd internal classification of the dinosaurs. He splits Saurischia up into three suborders, with the larger carnivores being more closely aligned with the prosauropods and sauropods than the small carnivores. This seems to follow Friedrich von Huene’s Pachypodosauria hypothesis, which you may remember from the earlier post about paleopods. This was the idea that the big “carnosaurs” are distinct from the small “coelurosaurs” and actually more closely related to the sauropodomorphs (making Marsh’s original Theropoda polyphyletic). While Wilfarth does indeed group all the small saurischians into Von Huene’s Coelurosauria, the group that Von Huene would have called Carnosauria is instead here referred to as… Theropoda. And includes the plateosaurid prosauropods, as per Marsh. My only guess is that he preferred Theropoda over Carnosauria based on priority rules, it being the older name.
He himself funnily enough notes how confusing all of these names get, though for a different reason. The name Theropoda, he correctly notes, makes no sense, because these dinosaurs had feet like birds, not mammals (Wilfarth 1949a, p. 20). Sauropoda is even worse, because the sauropods had the least lizard-like feet of all dinosaurs (Wilfarth 1949a, p. 34). This results in a tangent that I like to imagine many taxonomists would secretly agree with, even if some might not like to admit it:
“The unfitting and meaningless names have scared off many a friend of paleontology from dealing with the dinosaurs. The student can in the name of a saurian or a group of saurians not find anything about their characteristics. He therefore hardly remembers them and it is not easy for him to gain an overview. This unpleasant phenomenon is a consequence of the, in theory, useful and necessary law of priority. In my opinion, however, it would suffice if this was only valid for genera and species. The descriptors of superorders, orders, suborders and families would have to be determined, after enough clarity has been gained, through vote by experts, if obvious nonsense is not to be hitchhiked through the millennia.” (Wilfarth 1949a, p. 20 – 21, translated by me)
Despite their size and small heads, Wilfarth identifies prosauropods like Plateosaurus as carnivorous. Or rather faunivorous, which I have been told is the more useful term for predators feeding on small animals whole. Here he goes quite into detail on how he imagines Earth’s ecology at the time. Instead of land plants, the primary producers in almost all continental ecosystems must have been plankton and algae, which the tides pushed into the land and where they fed a wide variety of invertebrates (Wilfarth 1949a, p. 31). Here Wilfarth delves into the minutiae of all the different critters a tidal dinosaur could have fed on, presenting an extensive list of fauna which I suspect were inspired by personal visits to the German Wattenmeer. There must have been snails, clams, cephalopods, sea lilies, corals, anemones, jellyfish, brachiopods, infaunal crustaceans like the Schlickkrebs (Corophium) and a huge variety of worms, with Wilfarth (1949a, p. 29 – 31) naming 15 different genera of sessile tube worms that could have been on the menu. A dinosaur like Plateosaurus, with its small head and rake-like teeth, was perfectly suited to sieve through the sands and effectively “flesh-graze” on these rich invertebrate lawns (Wilfarth 1949a, p. 32), allowing it to grow so large.
Fig. 14: The watt worms which inhabit the intertidal zones. According to Wilfarth, such invertebrates must have made up the basis of most ecosystems (Source: Wilfarth 1949a, p. 42). |
Interesting to note here is that Wilfarth characterizes Plateosaurus as an obligate biped on land, on the basis that its arms were quite short and seemed unsuited for walking (Wilfarth 1949a, p. 33), substantiating this argument based on earlier statements made by Von Huene (Wilfarth 1938a, p. 269). Even in water, the arms were not used for locomotion but at best for anchoring the animal to the ground in case of strong currents. This is again notable, because until relatively recently Plateosaurus was commonly reconstructed as only a facultative biped, able to switch between a two- and four-legged stance. It took until Bonnan & Senter 2007 for it to be universally recognized that the animal could not pronate its wrists and therefore not use its hands for walking. So here we have another case of unexpected prescience baked into Wilfarth’s eccentricity.
This concludes Part 1. Be sure to check in again in a few days when we look at how Wilfarth interpreted the sauropods and theropods of the Jurassic and Cretaceous!
If you liked this and other articles, please consider supporting me on Patreon. I am thankful for any amount, even if it is just 1$, as it will help me at dedicating more time to this blog and related projects. Patrons also gain early access to the draft-versions of these posts and my art.
Related articles:
- The Alien Prehistoric World Trope - Part 2
- The weirdest things people have thought about pterosaurs
- Paleopods: The myth of the predatory prosauropods that kinda turned out to be true
- "Hawkinspunk" Spinosaurus
References:
- Benton, Michael: Scientific Methodologies in Collision. The History of the Study of the Extinction of the Dinosaurs, in: Evolutionary Biology, 24, 1990, p. 371-400.
- Bertozzo, Filippo; Manucci, Fabio; Dempsey, Matthew; Tanke, Darren; Evans, David; Ruffell, Alastair; Murphy, Eileen: Description and etiology of paleopathological lesions in the type specimen of Parasaurolophus walkeri (Dinosauria: Hadrosauridae), with proposed reconstructions of the nuchal ligament, in: Journal of Anatomy, 238, 2020.
- Bonnan, Matthew & Senter, Phil: Were the basal sauropodomorph dinosaurs Plateosaurus and Massospondylus habitual quadrupeds?, in: Barrett, P.M.; Batten, D.J. (eds.): Evolution and Palaeobiology of Early Sauropodomorph Dinosaurs (Special Papers in Palaeontology 77), Oxford 2007, p. 139–155.
- Byrne, H. M.; Green, J.; Balbus, S.; Ahlberg, P: Tides. A key environmental driver of the osteichthyan evolution and the fish-tetrapod transition?, in: Proc. R. Soc. A, 476, 2020.
- Colbert, Edwin: Review. Die Lebensweise der Dinosaurier, in: Journal of Paleontology, 24, 1950, p. 116.
- Colbert, Edwin: Relationships of the Saurischian Dinosaurs, in: American Museum Novitates, 2181, 1964, p. 1 – 24.
- Cuvier, Georges: Discours sur les Révolutions de la Surface de la Globe, et sur les Changements qu’elles ont Produites dans la Regne Animal, Paris 1825.
- Desmond, Adrian: The Hot-Blooded Dinosaurs. A revolution in Paleontology, London 1975.
- De Winter, Niels; Goderis, Steven; Van Malderen, Stijn; Sinnesael, Matthias; Vansteenberge, Stef, Snoeck, Christophe; Belza, Joke; Vanhaecke, Frank; Claeys, Philippe: Subdaily-Scale Chemical variability in a Torreites Sanchezi Rudist Shell: Implications for Rudist Paleobiology and the Cretaceous Day-Night Cycle, in: Paleooceanography and Paleoclimatology, 35, 2020.
- Ekman, Martin: A Concise History of the Theories of the Tides, Precession-Nutation and Polar Motion (From Antiquity to 1950), in: National Land Survey. Division of Geodetic Research, S-801, 1993, p. 585 – 617.
- Fulda, Ernst: Die Entstehung der Zechsteinsalze nach der Grossflutenhypothese von Martin Wilfarth, in: Kali, 32, 1937a.
- --1937b: Die Grossflutenhypothese und ihre Anwendbarkeit auf die Entstehung der Salzlagerstätten, in: Forschung und Fortschritt, 14.
- --1938a: Steinsalze und Kalisalze, in: Beyschlag-Krusch-Vogt: Die Lagerstätten, 3, 2, Stuttgart.
- --1938b: Salze, in: Geologische Jahresberichte, 1.
- --1939: The Theory of the Great Tides and its Application to the Theory of the Formation of the Salt Deposits, in: Research and Progress, 5.
- Galton, Peter: Ornithischian Dinosaurs and the Origin of Birds, in: Evolution, 24, 1970, p. 448 – 462.
- Hallett, Mark & Wedel, Mathew: The Sauropod Dinosaurs. Life in the Age of Giants, Baltimore 2016.
- Heilmann, Gerhard: The Origin of Birds, London 1926.
- Holland, William Jacob: The Skull of Diplodocus, in: Mem. Carnegie Mus., 9, 1924, S. 379 - 404.
- Kermack, Kenneth: A note on the habits of the sauropods, in: Annual Magazine of Natural History, 12, 1951, S. 830 – 832.
- Knoll, Fabien; Galton, Peter; Lopez-Antonanzas, Raquel: Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus, in: Geobios, 39, 2006, p. 215 – 221.
- Loomis, Frederic Brewster: Momentum in variation, in: The American Naturalist, 39, 1905, p. 839 – 843.
- Matthew, William Diller: Dinosaurs, New York 1915. (Readable here)
- McLoughlin, John: Archosauria. A New Look at the Old Dinosaur, New York 1979.
- McLoughlin, John: Synapsida. A New Look into the Origin of Mammals, New York 1980.
- McLoughlin, John: The Tree of Animal Life. A Tale of Changing Forms and Fortunes, New York 1981.
- Naish, Darren: The response to and rejection of Brian Ford’s Too Big to Walk, a 21st century effort to reinstate the aquatic dinosaur hypothesis, in: Historical Biology, 2024.
- Nölke, Friedrich: Kann dem Monde ein Einfluss auf die geologische Entwicklung eingeräumt werden?, in: Bremer Beiträge zur Naturwissenschaft, 1935.
- Norman, David: The Illustrated Encyclopedia of Dinosaurs, London 1985.
- Paul, Gregory Scott: Predatory Dinosaurs of the World. A Complete Illustrated Guide, New York 1988.
- Romer, Alfred Sherwood: Osteology of the reptiles, Chicago 1956.
- Seeley, Harry Govier: On the Classification of Fossil Animals commonly named Dinosauria, in: Proceedings of the Royal Society, 43, 1887, p. 165–171.
- Sternberg, Charles Mortram: Were there Proboscis-bearing Dinosaurs? Discussion of Cranial Protuberances in the Hadrosauridae, in: Annals and Magazine of Natural History, 11, 1939, p. 556 – 560.
- Vidal, D.; Mocho, P.; Aberasturi, A.; Sanz, J. L.; Ortega, F.: High browsing skeletal adaptations in Spinophorosaurus reveal an evolutionary innovation in sauropod dinosaurs, in: Scientific Reports, 10, 2020.
- Wiersma, Kayleigh & Sander, Martin: The dentition of a well-preserved specimen of Camarasaurus sp.: implications for function, tooth replacement, soft part reconstruction, and food intake, in: Paläontologische Zeitschrift, 91, 2017, p. 145 – 161.
- Wilfarth, Martin: Sedimentationsprobleme in der Germanischen Senke zur Perm- und Triaszeit, in: Geologische Rundschau, 24, 1933, p. 349 – 377.
- --1934: Strömungserscheinungen im Wellenkalkmeer, in: Zeitrschrift der Deutschen Geologischen Gesellschaft, 86, p. 265 – 285.
- --1936: Die Gezeiten im Meere des Malm Zeta bei Solnhofen, in: Zeitschrift der Deutschen Geologischen Gesellschaft, 88, p. 57 – 61.
- --1938a: Die Sauropoden als Bewohner des Grossgezeitenraumes, in: Paläontologische Zeitschrift, 19.
- --1938b: Was hat die Grossgezeitenhypothese zum Problem der Erdölentstehung zu sagen?, in: Kali, 32, 1938b, H11.
- --1938c: Kalkfällung im Wellenkalkmeer, in: Beiträge des Geologischen Thüringischen Vereins, 5, p. 46 – 48.
- --1938d: Gab es rüsseltragende Dinosaurier?, in: Zeitschrift der Deutschen Geologischen Gesellschaft, 90, p. 88 – 100.
- --1939: Die Nasenbasis der Lambeosaurinae, in: Zentr. Bl. F. Min., 1, p. 24 – 39.
- --1940a: Der Atemrüssel der Hadrosauriden, Halle.
- --1940b: Die Umdrehung der Wirbeltierahnen, Halle.
- --1947: Rüsseltragende Dinosaurier, in: Orion, 2, 11/12, p. 525.
- --1948: Grossgezeiten in der Erdvergangenheit, in: Orion, 3, H 2/3, p. 79.
- --1949a: Die Lebensweise der Dinosaurier, Stuttgart.
- --1949b: Leben heute noch Saurier?, in: Prisma. Illustrierte Monatsschrift für Natur, Forschung und Technik, 4, 6.
- Wiman, C.: Über einige neue Lebendbilder von Dinosauriern, in: Paläontologische Zeitschrift 23, 1942, p. 237 – 249.
- Witmer, Lawrence: Nostril Position in Dinosaurs and Other Vertebrates and Its Significance for Nasal Function, in: Science, 293, 2001, 850 – 853.
Absolutely spectacular research. I had no idea any of this existed. Thank you so much for bringing it to light. I look forward to seeing what you have to say in your future posts.
ReplyDeleteWow! I heard about the trunked hadrosaurs from Sternberg's paper *years* ago, but was never ever to find the cited study or associated art. It's been at the back of my mind since then, and here I find a tell-all about it! Very happy to see this, and that the story itself is more interesting than Sternberg's fiction ever was. Can't wait for the following parts.
ReplyDelete