Deinosuchus: the Dalek-backed alligatoroid that (sometimes) made chew toys of dinosaurs

Deinosuchus rugosus swallows the remains of a large Cretaceous sea turtle. Other archosaurs notice, decide to interrupt.

Any palaeontological geek worth their salt knows that several gigantic crocodyliform species - colloquially called 'supercrocs' - have appeared in the last 100 million years. They include the Moroccan, Cretaceous pholidosaurid Sarcosuchus imperator, several species of the South American Miocene caimanine Purusaurus, and the grandfather of them all, Deinosuchus, from the Late Cretaceous (Campanian) of North America. Deinosuchus is not the most 'extreme' of these giant crocodyliforms in terms of anatomy or size (but see below), but discovery of a partial skeleton in 1903 (and description six years later) was the first evidence for some croc-line archosaurs being very, very large. It is also perhaps the best publicly known giant crocodyliform, various reconstructions of its skull and skeleton appearing in museums all over the world, and being a semi-regular component of palaeoart.

Perhaps because Deinosuchus is 'only' a giant crocodylian and not a member of a completely extinct, weirdo lineage, coverage of its palaeobiology is often limited to factoids on its immense size and probable habits of eating dinosaurs. Our short attention span for this animal is not new: one of the key players in its discovery, John Bell Hatcher, lost interest in describing the first significant Deinosuchus remains once its crocodylian identity (rather than dinosaurian, as originally supposed) became apparent in 1903 (Schwimmer 2002). It took further persuasion and a number of years for palaeontologists to actually publish, name and describe this animal after Hatcher died in 1904 (Holland 1909). As we'll discover in this article, the scant attention paid to this animal is rather criminal: over the last century a detailed and fascinating picture of Deinosuchus has developed.

The Colbert and Bird (1954) reconstruction of Deinosuchus riograndensis, based on partial skull material collected from the late Campanian Aguja Formation, Texas. We now appreciate this reconstruction as being erroneous in a number of crucial ways, and it should not be considered representative of the appearance of this animal. Note the apertures in the snout tip alongside the actual nares. Images from Colbert and Bird (1954).

I'm going to start by sidestepping the confused history surrounding the discovery and naming of Deinosuchus material - interested parties can find a full summary in David Schwimmer's (2002) book Deinosuchus: King of the Crocodylians. It will suffice to state that fossils of this animal have been known since at least the mid-1800s and repeatedly hopped between species and genera during the last 150 years. Only one or two species are recognised nowadays. D. rugosus is the type species of the genus, first identified from two characteristically large, blunt teeth with wrinkled, thick enamel collected from the eastern US in the 1858 (examples below). Over time, these teeth were found to be linked with other Deinosuchus material including extremely thick, massive and deeply pitted osteoderms. These elements are highly characteristic even today, and permit even isolated teeth and osteoderms to be referred to this species (Schwimmer 2002). A possible second species is D. riograndensis, based on very large Deinosuchus fossils from Texas recovered in the 1940s (Colbert and Bird 1954). It's this riograndensis material that most of us think of when Deinosuchus is mentioned, it providing the basis of a famous, 2 m long and largely artistic Deinosuchus skull reconstruction unveiled at the AMNH in 1954 (above). The riograndensis skull was once thought characteristic because of unusual openings in the side of the snout tip (Colbert and Bird 1954), but these are now thought to be damage caused during preparation (Schwimmer 2002). As more and better Deinosuchus remains have been recovered in recent decades, some have argued riograndensis and rugosus are one and the same animal, the latter holding nomenclatural priority. Full agreement on this does not seem apparent from current literature, but on-going work on relatively complete Deinosuchus material will help clarify the taxonomy of this animal in future.

Deinosuchus is recognisable from its teeth alone. These specimens are from the posterior end of the jaws, and show the wrinkled enamel typical of the genus. Note the heavily worn and broken the tips. From Schwimmer (2010).

We now appreciate that Deinosuchus is one of the oldest members of Crocodylia, the crown group crocodile-line archosaurs. Features of its skull and jaws indicate it was specifically a member of Alligatoroidea, the lineage of Crocodylia represented today by alligators and caimen. It's not uncommon to see Deinosuchus referred to as a 'giant alligator', but this is not really accurate and I recommend avoiding such terminology, even for lay audiences. Alligators and Alligatoridae are a distinctive group of alligatoroids with particular habits and anatomy, and they are no more closely related to Deinosuchus than caimen. It must also be stressed that neither of our modern alligatoroid clades are especially closely related to Deinosuchus. I figure most people will intuitively grasp the rough meaning of 'giant alligatoroid', and suggest this term is used in preference of 'alligator' in outreach media about this animal.

The Deinosuchus fossil record is something of a mixed bag. There are hundreds of fossils of it, but most of them are isolated postcranial bones, broken bits of skull and, especially, those massive teeth and osteoderms. The state of many Deinosuchus fossils can be ascribed to its remains being reworked by storms after their initial burial. Some partial skeletons and more complete skulls escaped this treatment but are not yet described in detail. A silver lining to not having much in the way of complete material is that isolated Deinosuchus bones are distinctive enough to map its range across Campanian North America. Many of us might think of Deinosuchus as a Texan animal, but it actually enjoyed a wide distribution, and being most abundant in the southeastern United States. A clear palaeobiogeographical pattern can be gleaned from Deinosuchus fossils, a divide separating occurrences in Montana, Wyoming, Utah, Colorado, New Mexico, Texas, and Northern Mexico from remains on the eastern side of the United States - Mississippi, Alabama, Georgia, North Carolina, and New Jersey (Titus et al. 2008). This east-west distribution is no fluke of preservation but reflection of Deinosuchus populations being separated by the Western Interior Seaway, the continental sea which divided North America during the Cretaceous. Although apparently not a fully marine creature, it is thought Deinosuchus lived in the coastal waters and estuaries of this seaway as, to date, its fossils have not occurred in fully freshwater or terrestrial deposits. Further evidence of its preference for coastal waters is the recovery of more complete and associated remains from wholly marine deposits.

A modern depiction of the Deinosuchus rugosus skull and mandible. From Schwimmer (2002).

Our incomplete knowledge of Deinosuchus anatomy means we can only form a partial picture of what it looked like in life. We can say, however, that the famous 'Colbert and Bird' riograndensis skull sculpture from the 1950s is erroneous in several regards. Manufactured as a display for the American Museum of Natural History (and then reproduced for museums around the world), the Colbert and Bird skull is too large, the snout too narrow, the tooth morphology inexact and, as noted above, the openings in the snout tip are likely erroneous (Schwimmer 2002). This is not a dig at AMNH artists course: they did the best they had with material available to them, and it's only with the discovery of better fossils that we can now spot errors. Perhaps unfortunately, museums continue to display this reconstruction and artists continue to use it as a reference. A modern picture of Deinosuchus is rather different: a broad- and deep-snouted crocodylian with a skull known to be at least 1.3 m in length, and perhaps a little longer if very fragmentary remains are being correctly interpreted (Schwimmer 2002). Most of its cranial features are typically crocodylian, including the presence of huge spaces for jaw muscle attachment, development of a secondary palate and dorsally situated orbits and nares. Befittingly for such an enormous animal, the teeth are huge and consistently robust along the jaw, those at the front being conical and pointed, and those at the back being increasingly stunted and shortened. The toothrow is very long in stretching from the jaw tip to just behind the eye. All teeth have the distinctively thickened, wrinkled enamel mentioned above. Unlike modern alligatoroids, a notch in the side of the upper jaw acts as a receptacle for the fourth tooth of the lower jaw and presumably rendering it visible even when the jaw was closed. This is the 'primitive' condition for crocodylians, and means that although Deinosuchus is an alligatoroid, it likely had a crocodile's smile.

Less can be said about the body and limbs of Deinosuchus. Schwimmer (2002) reports that partial skeletons hint at a general form and proportion not unlike a modern alligator. However, some authors have noted discrepancies in scaling of Deinosuchus limb bones which might indicate reduced limbs in at least the largest specimens (Farlow et al. 2005, see below). One thing we can be sure of is that the body of Deinosuchus was covered in those aforementioned large, thickened osteoderms (below). The exact arrangement of these elements remains unknown, but we can predict that at least four rows of osteoderms extended along the body of Deinosuchus because of its affinities to modern crocodylians. These osteoderms become disproportionately massive and robust with growth, so that those of the largest individuals are distinctively chunky and have lost some definition of a keel found in smaller examples. Artists should take note of this: the dorsum of a big Deinosuchus would have looked more like a gnarly Dalek chassis than the back of any modern crocodylian. As is typical for crocodyliforms, these dermal bones might have reinforced the trunk skeleton as well as providing armour plating, forming a network of muscle, ligaments and bone which bound the torso together (Salisbury and Frey 2000). It is speculated that the presence of very large, robust osteoderms in the biggest Deinosuchus indicates the presence of a torso strong enough for terrestrial locomotion (Schwimmer 2002).

The huge, deeply pitted and bulbous scutes which characterise Deinosuchus, as illustrated by Holland (1909). This image is a composite of two scutes from Holland's work, put together by FanCollector for Wikipedia. Both show cervical osteoderms, the left being a particularly big one

The maximum size of Deinosuchus is the source of much fascination and discussion. The largest estimates based on reliably measured remains suggest body lengths of around 12 m (Schwimmer 2002), but - as usual with giant fossil animals - there are a number of factors and caveats worth considering here. The scant nature of Deinosuchus fossils dictates that we must extrapolate the size of large Deinosuchus from much smaller, better known individuals and modern crocodylians. The 12 m figure stems from scaling the largest Deinosuchus vertebrae to total body length estimates of smaller individuals (Schwimmer 2002). Skulls and mandible lengths, when compared to a dataset of modern alligator proportions, indicate the largest animals achieved 10 m in length (Schwimmer 2002; Farlow et al. 2005). Another approach, using femoral measurements, results in a maximum body length estimate of 6-8 m, this being estimated from big femora typically thought to indicate 10 m+ animals. The explanation for these differences will be familiar to anyone who's estimated the size of a big, extinct animal: uncertainty about the proportions of the animal in question, the need to extrapolate well beyond the size boundaries of modern analogues, and the lack of associated remains of the biggest individuals. Most workers seem happy with a 10 m length estimate for a big Deinosuchus, those lower estimates based on femoral size being explained as possible evidence of reduced limb proportions in the biggest Deinosuchus individuals (Farlow et al. 2005). As usual, we await the discovery of more complete and informative remains to tell us the full story here. It should be stressed that similar caveats apply to size estimates of other 'supercrocs': despite the media hype associated with some discoveries, it's quite difficult to know which crocodyliform species was largest based on our current material.

Regardless of what the actual maximum size of Deinosuchus was, we have good reason to think that many individuals were not true giants. No specimens indicative of 10 m body length have been found among the many hundreds of Deinosuchus remains from the eastern side of the US, it being instead thought that eastern Deinosuchus didn't grow longer than 8 m. That's still pretty big of course, but not too far off crocodylians that we're familiar with today (the biggest saltwater and Orinoco crocodiles on record are a little over 6.5 m - Grigg and Krishner 2015). The true giants only occur in the west, and are much rarer fossils than their eastern counterparts. These fossils are also slightly younger than the eastern specimens, perhaps indicating changes across time and geography were responsible for Deinosuchus becoming exceptionally large. Research into the growth rates of Deinosuchus indicate that there might be nothing unusual about it's growth trajectory despite its size. It seems to have grown with a similar strategy to other crocodylians - relatively fast at first, and progressively slower over time - but simply stretched out the growth duration to many decades (Erickson and Brochu 1999). Growth rings in osteoderms indicate that the largest animals were about around 50 years old (below).

Growth rates in living and extinct crocodylians (a) and growth rings in a Deinosuchus osteoderm (b). Note how the Deinosuchus growth trajectory is essentially a scaled up version of its smaller relatives. From Erickson and Brochu (1999).

For many, the main discussion to be had about Deinosuchus is the impact it had on local dinosaur populations: was this animal a dinosaur predator? Attempts to answer this question stem from two sources: biomechanics and fossil evidence of ancient faunal interactions. The first, biomechanics, includes a recent study of 'death rolling' (the crocodylian habit of rotating around the long axis of the body while gripping prey with their jaws, literally twisting it apart) and whether Deinosuchus could use this strategy to dismember large prey (Blanco et al. 2015). Snout strength can be correlated quite accurately to death rolling capabilities in modern crocodylians (Blanco et al. 2015) and, seeing as this can be inferred from upper jaw skeletons alone, we can obtain some insight into the death rolling capabilities of extinct crocodyliforms. Perhaps surprisingly, estimates of Deinosuchus jaw strength were approximately one third of the strength required for this behaviour (Blanco et al. 2015). Scaling factors have been used to explain this unexpected result and, despite the outcome of their experiments, Blanco et al. suggest that death rolling was possible in Deinosuchus. I must admit to thinking additional experimentation is needed to quantify those scaling factors before considering this matter closed. Moreover, in checking the Blanco et al. data for this article I noted that their study modelled the Deinosuchus skull as 1.8 m long, a figure noted as speculative by Schwimmer (2002) and almost 40% longer than the largest measured skull length reported by the same author (1.31 m). A 40% shorter skull would be less prone to the scaling effects outlined by Blanco et al. and may result in a jaw strength more suited to death rolling - it would be great to see this checked out in future.

Partial theropod hindlimb bone (tibia or metatarsal) post a one-on-one session with Deinosuchus jaws. This bone is meant to be subrounded in cross section. From Schwimmer (2010).

More positive and definitive answer about Deinosuchus dinosaur predation stems from fossil evidence. It seems that, yes, Deinosuchus did dine on dinosaurs, but not exclusively or maybe even often (Schwimmer 2010). A handful of Campanian dinosaur bones - including a theropod hindlimb element from Georgia (above) and hadrosaur vertebrae from Texas - possess bitemarks characteristic of crocodylians, albeit on a scale unseen in the modern day (Schwimmer 2002, 2010). The theropod bone can only be described as exceptionally chewed: numerous, overlapping circular potmarks show where the bone was repeatedly bitten and crushed by a very powerfully jawed animal. So pulverized is this bone that its once subrounded cross section has become quadrangular - Schwimmer (2002) summarises the state of this specimen as 'resembling a dog's worn chew toy' (p. 186). The crocodylian signature, bite mark size and provenance of both specimens point to Deinosuchus as a possible perpetrator, and evidence that it did eat dinosaurs on occasion. Of course, it cannot be easily established whether these animals were killed by Deinosuchus or merely scavenged by them - some reasoning for the former is discussed by Schwimmer (2010).

Deinosuchus bite marks in fragments of a turtle (Chedighaii barberi) plastron. The tooth marks are about 4-5 times larger than those made by 4 m long nile crocodiles. From Schwimmer (2010).

There are reasons to think dinosaur meat was not a mainstay of Deinosuchus diet, however. Whereas a few dinosaur bones have been linked to the jaws of this crocodylian, more than a dozen Campanian sea turtle specimens have been found with bite marks made by a giant crocodylian (Schwimmer 2010). A diet of turtles is not surprising when we consider that the skull and dentition of Deinosuchus is more adapted to crushing bone than piercing skin and flesh (Schwimmer 2002): those robust posterior teeth are especially reminiscent of teeth in modern, turtle-eating crocodylians. Many Deinosuchus teeth are considerably worn and broken too, a likely consequence of being smashed into hard, bony prey rather than soft, spongy dinosaur limbs. Healed turtle shells suggest that these animals were predated by Deinosuchus rather than just scavenged, and raking bites across some specimens may record less fortunate turtles being juggled about Deinosuchus jaws when being eaten (Schwimmer 2010). The coastal and estuarine environmental bias of Deinosuchus fossils is consistent with it being a serial turtle predator, this being ideal habitat to find sea-going prey. Curiously, other marine inhabitants of the Western Interior Seaway have yet to be associated with Deinosuchus bite marks: perhaps it really did have a preference for turtles, or perhaps other skeletons were simply pulverised beyond recognition by those massive jaws. Either way, our discussion of this animal's feeding habits would not be complete without mentioning the numerous, possible Deinosuchus coprolites which have recently been identified (Harrell and Schwimmer 2010). Sadly, these do not reveal much about diet or digestive anatomy, other than the obvious fact that Deinosuchus poop was on average a lot larger than that produced by other crocs. Several anomalous features of these coprolites have led some authors (e.g. Hunt and Lucas 2010) to be sceptical of their organic origins however, their alternative being that they are simply calcareous nodules.

The foraging habits of Deinosuchus brings us to new perspectives on where it fits into Mesozoic ecology. New evidence is eroding the uniqueness of Deinosuchus in Campanian North America, it no longer being the only very large or even giant crocodyliform species in some localities. These new finds include a currently poorly known, but obviously giant neosuchian from the Williams Fork Formation of Colorado (Foster and Hunt-Foster 2015) and a more completely known, 7 m long, undescribed neosuchian from Woodbine, Texas (Main 2012). The latter is currently being worked on, and early indications are that it might represent a late surviving goniopholidid - a much older branch of the crocodyliform lineage. Whatever they turn out to be, the Woodbine and Williams Fork animals suggest that very large crocodyliforms might not have been unusual in Campanian North America. Their presence in a timeframe deficient of large theropods has not gone unnoticed, it being speculated that these large crocodyliforms may have been doing work normally reserved for big predatory dinosaurs (e.g. Schwimmer 2002). Similar proposals have been made about other large bodied Late Cretaceous carnivores taking over typically theropodan roles (e.g. Witton and Naish 2015) - the notion of the Mesozoic as an all-dinosaur show is looking increasingly out of date.

As a closing thought, I find it interesting that we tend to portray Deinosuchus as something of a freak species, one of those rare forays of crocodylian evolution into gigantic size which never really seemed to last that long or lead anywhere. As might be apparent from this article, this view is somewhat misleading. Deinosuchus certainly represents an 'extreme' of crocodylian evolution, but it's at the end of a spectrum, not a weird outlier from the rest of the group. Much of what it did, how it did it, and what makes it a fascinating animal, is mirrored in its modern and fossil relatives. Contrary to some perspectives on this animal, the fact it represents an ancient member of a modern group does not make it tedious or dull. Quite the opposite is true: Deinosuchus reminds us that animals from Deep Time are part of a continuum with our own fauna, revealing the awesome things modern lineages have been capable of, the potential their anatomies have in the present, and what they might be up to in future. How anyone can find pondering an animal that gives such a raw perspective on evolution and adaptation boring or uninteresting is beyond me.

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References

• Blanco, R. E., Jones, W. W., & Villamil, J. (2015). The ‘death roll’of giant fossil crocodyliforms (Crocodylomorpha: Neosuchia): allometric and skull strength analysis. Historical Biology, 27(5), 514-524.
• Colbert, E. H., Bird, R. T., & Brown, B. (1954). A gigantic crocodile from the Upper Cretaceous beds of Texas. American Museum Novitates; no. 1688.
• Erickson, G. M., & Brochu, C. A. (1999). How the ‘terror crocodile’ grew so big. Nature, 398, 205-206.
• Farlow, J. O., Hurlburt, G. R., Elsey, R. M., Britton, A. R., & Langston Jr, W. (2005). Femoral dimensions and body size of Alligator mississippiensis: estimating the size of extinct mesoeucrocodylians. Journal of Vertebrate Paleontology, 25(2), 354-369.
• Foster, J. R., & Hunt-Foster, R. K. (2015). First report of a giant neosuchian (Crocodyliformes) in the Williams Fork Formation (Upper Cretaceous: Campanian) of Colorado. Cretaceous Research, 55, 66-73.
• Grigg, G., & Kirshner, D. (2015). Biology and Evolution of Crocodylians. Csiro Publishing.
• Harrell, S. D., & Schwimmer, D. R. (2010). Coprolites of Deinosuchus and other crocodylians from the Upper Cretaceous of western Georgia, USA. New Mexico Museum of Natural History and Science, Bulletin, 51, 209-213.
• Holland, W. J. (1909) Deinosuchus hatcheri, a new genus and species of crocodile from the Judith River beds of Montana. Annals of the Carnegie Museum, 6, 281–294.
• Main, D. J. (2012). Crocodiles of the Texas Cretaceous; the Campanian of Big Bend to the Cenomanian of North Texas, a comparison of great size, feeding behaviour and paleoecology. Geological Society of America Abstracts with Programs, 44, 3.
• Schwimmer, D. R. (2002). King of the crocodylians: the paleobiology of Deinosuchus. Indiana University Press.
• Schwimmer, D. R. (2010). Bite marks of the giant crocodylian Deinosuchus on Late Cretaceous (Campanian) bones. New Mexico Museum of Natural History and Science Bulletin, 51, 183-190.
• Titus, A.L., Knell, M.J., Wiersma, J.P., Getty, M.A. (2008). First report of the hyper-giant Cretaceous crocodylian Deinosuchus from Utah. Geological Society of America Abstracts with Programs, 40, 58.
• Witton, M. P. and Naish, D. (2015) Azhdarchid pterosaurs: water-trawling pelican mimics or "terrestrial stalkers"? Acta Palaeontologica Polonica 60, 651-660.

New takes on the Wealden Supergroup palaeobiota, part 2: Baryonyx, freshwater plesiosaurs, ornithomimosaurs and others

Last week we took a look at some new art of animals from the Wealden Supergroup, the intensively studied, historically important Lower Cretaceous rocks of Southern Britain. We all know the Wealden for celebrity dinosaurs like Iguanodon and Baryonyx, but there's a heap of other interesting animals in there which get relatively little publicity. It's mostly these we're focusing on here, in the second (and final) part of these 'picture of the day'-style posts. 

As before, if you like anything here, remember that you can buy prints of them all from my shop (the Wealden section might be relevant) and its new Facebook outlet. Indeed, if you like my work and are on Facebook, why not 'like' the new Mark Witton Palaeoart page? It's the best place to see when new prints and finished pictures are available.

Baryonyx walkeri: king of the fishers, redux

Baryonyx walkeri, off for a stroll among the crocodyliforms and pterosaurs.

Let's break this post in with a familiar animal: spinosaurid Baryonyx. It's hard to appreciate now how weird this animal seemed back in the 1980s and 1990s. At this point, other spinosaur material was only very poorly known, and laymen and scientists alike found this weird, superficially-crocodile like animal fascinating. Ironically, it's recently turned out that we first collected Wealden spinosaur material centuries ago, but struggled to recognise its significance until more complete remains were unearthed in the 1980s. We now know that Baryonyx can be found throughout a good chunk of upper Wealden stratigraphy and teeth referable to it - or another spinosaurid - are fairly common, at least as Wealden dinosaur fossils go. Baryonyx provided the basic template we'd recognise for all spinosaurid anatomy until last year when, famously, some spinosaurs were proposed to be rather different. It's clear that, whatever is going on with Spinosaurus, Baryonyx retains more conventional hindlimb and pelvic proportions, and may not have been so aquatically adapted as true spinosaurines. In this updated image, B. walkeri is splashing into a body of water while goniopholidid crocodyliforms and gnathosaurine pterosaurs go about their business around it. Note how much larger Baryonyx is compared to the crocs: Baryonyx is the largest theropod in the Wealden Supergroup, by a good margin.

Button-toothed crocs, redux

Bernissartid Koumpiodontosuchus aprosdokiti foraging for molluscs. It's eating a mud snail, Viviparus cariniferus, while tiny (6 mm long) physid gastropods Prophysa crawl over pond scum in the lower left of the image. Dragonflies provide scale, and unnamed tetanurans prowl around the background.

Last year I was lucky enough to provide the first restoration of Kompiodontosuchus aprosdokiti, a small neosuchian crocodyliform common to the Wessex Formation, and perhaps other parts of the Wealden sequence. Koumpiodontosuchus is a bernissartid, a group of small-bodied crocodyliforms with robust, shell-cracking teeth at the back of their jaws. As you'll know if you read my write up last year, these were likely employed in smashing molluscs and insects. The tetanuran theropods in this image are unnamed, but are not thought to be referable to any existing Wealden taxa. We probably need more material of them to consider them nameable, however: recognising that they are different from other Wealden theropods is only half the battle. Modern students of Wealden fossils famously do their best to preserve historic names based on fragmentary bones, but there seems to be an effort to 'future proof' Wealden taxonomy against confusion by only naming well-represented, characteristic animals. I guess I could have chosen one of the better known theropods to play the 'This was the Age of Dinosaurs' card for this PR image, but I think it's good to show that not all large theropods in the Wessex palaeobiota were Neovenator, Baryonyx or Eotyrannus. 

Welcoming the new Wealden ornithomimosaurs

A flock of Wessex Formation ornithomimosaurs forage in a marshland, while istiodactylid pterosaurs skulk about behind them.

Those keeping their ears to the ground will know that the newest arrivals to the Wealden dinosaur palaeobiota are ornithomimosaurs, commonly known as ostrich dinosaurs. Two specimens show that these animals were present in both the Weald and Wessex basins of the broader Wealden succession, and one of these fossils represents a historic taxon named in 1889: Valdoraptor oweni. Key to identifying ostrich dinosaurs in the Wealden was the discovery of abundant ornithomimosaur remains in France, many of which are so reminiscent of Valdoraptor and other Wealden theropod material that they may represent the same taxon. If you want to know more about these and their relationship to the complex story of Wealden theropods, check out Darren Naish's post on this at Tetrapod Zoology.

The above new painting shows a group of (nameless) Wessex Formation ornithomimosaurs in a well-vegetated marshland, in the rainy season, while istiodactylid pterosaurs mosey about in the background. The abundance of ostrich dinosaurs and juveniles in the middle-right are nods to the frequent recovery of abundant specimens of different levels of maturity at many ostrich dinosaur sites, including the new, French 'Angeac ornithomimosaur'. Note that the wings of the running foreround animal are somewhat swept back: I don't think the more common way of reconstructing ornithomimosaurs with 'dangly arms' looks right. They look like they should be holding shopping bags or something.

Valdosaurus in the forest, redux

Two Wealden dryosaurids Valdosaurus canaliculatus, and a stubborn avialan.

Ornithomimosaurs weren't the only fast runners in Wealden landscapes. Dryosaurids, like Valdosaurus canaliculatus were also fleet-footed animals with powerful, well-muscled hindlimbs, and tiny bodies attached to the front. In this reworked image, two of these 3-4 m long animals are taking it slow through a Wealden woodland. Although Wealden climates were quite warm and arid, leaving much of the landscape looking quite chaparral-like, some relatively upland parts seem to have been more vegetated: it's here that this picture is set. In my mind, these animals always walked with the stooping posture of the foreground animal - as noted last time, I like the idea that prehistoric animals had characteristic postures varying slightly from those we consistently restore in skeletal restorations. Note the avialan on the left of the image, which is a nod to the recovery of bird teeth from Wealden deposits. Anyone who's ever been forced to walk around a stubborn reclined mallard will recognise the situation now facing the Valdosaurus.

Barilium dawsomi in leathers, redux

Barilium dawsoni, a large and very robust iguanodont from Sussex. A flock of 'Ashdown maniraptorans' add scale.

Last time we featured Iguanodon bernissartensis: now it's the turn of the 'other' big Wealden iguanodont, the stratigraphically older, and osteologically chunkier Barilium dawsoni. In this redone painting, I've tried to make the Barilium skin more interesting than just plain old scales, covering the back in small, horny ossicles and creasing the flanks as if the skin is particularly thick, leathery and folded. I think we should be rendering more interesting skin regularly in scaly dinosaur palaeoart, as it seems most extensive dinosaur skin remains show unexpected features - strangle scales, wattles, folds and that sort of thing - which small skin patches mostly cannot record adequately. It's interesting to contrast these skin impressions with homogeneous restorations of scaly dinosaur appearance presented by some, where every species is covered in smooth hide following perfect contours of the underlying tissues: I'm not sure that's what fossils are telling us. As before, the 'Ashdown maniraptoran' provides scale to the bulk of Barilium. For the uninitiated, the Ashdown maniraptoran is seriously small for a Mesozoic dinosaur - maybe about 30-50 cm long. If you find big iguanodonts exciting, be sure to check out this previous post.

Polacanthus redux, again

A Wealden tree vies for attention with Polacanthus foxii, and some tiny birds.

OK, I'm cheating a bit with this one. This redone version of a much older painting has been posted fairly recently, but it seemed a bit remiss to skip this ankylosaur in this run down of recently produced Wealden palaeoart. Polacanthus foxii is, of course, the Wealden's sacral-shield-bearing nodosaurid, shown here strolling around a Cretaceous hillock with some birds for company. Having scratched the completist itch, let's move on, because we've seen this all before.

Accidentally sinister Leptocleidus, redux 

Mother and calf Leptocleidus superstes, a freshwater leptocleidid plesiosaur, explore a river inlet in Lower Cretaceous Sussex. 

Our final stop is in Wealden rivers and estuaries, where Leptocleidus superstes and other species of freshwater leptocleidid plesiosaurs roamed. The new version of this image has added a lot of detail on top of the original, which has inadvertently made the mother and calf Leptocleidus look more sinister than intended - hey, it's not my fault their teeth stick out like that. Back in the original post on these animals I mentioned that pliosaurs may also have been present in Wealden lakes and rivers, but note that this is no longer certain: the Hastanectes valdensis remains once provisionally considered pliosauroid have been placed in Leptocleididae in more recent analyses. That does make for a neater story - it means that leptocleidids retain their dominant role as 'near-shore-or-freshwater' animals, but perhaps a slightly less interesting one.

And that's all for now - I hope you've enjoyed this jaunt back to the ancient Wealden and these revised artworks. I'm sure we'll visit the Wealden again in time. Coming next, probably: walking with non-pterodactyloid pterosaurs.

New takes on the Wealden Supergroup palaeobiota, part 1: Iguanodon, Neovenator, Eotyrannus and others

Regular readers will know that I'm prone to dabbling in palaeoart depicting the environments and animals of the Wealden Supergroup, the 18 million year stretch of Early Cretaceous time represented by mud-and sandstone deposits across the southern UK. Recently, I've been updating some existing Wealden work as well as producing some new stuff of other Wealden species. With no time to produce a new post of substance, here's a bumper 'picture of the day'-type post. Initially, I was going to chuck something like ten images on here, but time has run short and I'll have to split it in two.

If you like anything here, remember that you can buy prints of them all from my shop (there's now a Wealden section, too), which is now also browsable from the comfort of Facebook. OK, enough preamble: into the Wealden once again...

Iguanodon bernissartensis: thumb wars

Two Iguanodon bernissartensis, the quintessential Wealden iguanodont, decide to settle their differences, while members of their herd watch on.

Poor old Iguanodon doesn't get the attention it used to, and a lot palaeoart we do see of it tends to focus on tried and tested behaviours: lots of standing about and eating, but not much else. In this new painting, I've attempted to show two big Iguanodon individuals settling an intra-specific dispute via use of thumb spikes. Long-term readers may recall that we've covered iguanodont thumb spikes before, and that I. bernissartensis has especially big ones. Here, they've been swinging their thumbs at each other's soft bits, causing deep, bloody wounds. This might seem extreme, but there are plenty of modern animals which take intraspecific fights to similarly gory levels - elephant seals were a key inspiration here. I imagine battling Iguanodon would look like an armed sumo-wrestling match, albeit with longer tails and less rice. Note that you can see the breath of several animals here: Wealden winters are not meant to be especially warm.

Rebbachisaurids vs. Neovenator salerii redux

Carcharodontosaurian Neovenator salerii stalks a pair of rebbachisaurid sauropods, using darkness as cover.

A while back I posted about dinosaur predation, noting that modern animal predator acts are often far less gladiatorial and epic than we might imagine. It's this slow, considered approach to predation which I'm attempting to show here, as the carcharodontosaur Neovenator stalks two rebbachisaurid sauropods in the dead of night. The idea is that the Neovenator has much better eyesight than the sauropods, who know they're in trouble, but can't really respond adequately. Note the rain: some recent models of Wealden palaeoclimates suggest it was wetter than previously modelled (albeit with very high evaporation rates for much of the year).

Anteophthalmosuchus hooleyi vs. Hypsilophodon foxii, redux

Large goniopholidid Anteophthalmosuchus hooleyi takes advantage of a flooding river to hunt two stranded Hypsilophodon foxii.

Speaking of rain, we know that some parts of the Wealden were prone to flooding following particularly intense downpours. That's good news for animals adapted for powerful swimming, but less welcome to species which prefer dry land. Here, in this reworked painting, the large Wealden goniopholidid Anteophthalmosuchus hooleyi has found a stranded pair of adult and juvenile Hypsilophodon foxii, and is taking full advantage of the situation. Goniopholodids are a group of almost-crocodiles characterised by long forelimbs, interlocking scutes and overbitten jaws - you can read more about them here.

Eotyrannus lengi: firestarter, redux 

Early tyrannosauroid Eotyrannus lengi stalks the edge of such a wildfire. 

What else does rain bring? Sometimes, lightning. When introduced to a parched Wealden landscape, lightning strikes caused short-lived canopy fires which, ultimately, created conditions ideal for fossil preservation. In this reworked painting, a fully-feathered tyrannosauroid Eotyrannus lengi is prowling the periphery of a Wealden canopy fire to grab any animals flushed out by the flames.

The tiny wars of Wesserpeton evansae, redux 

Two Wesserpeton evansae get in each other's faces, because some animals are just jerks.

OK, enough about Wealden weather. Here's a reworked version of two of the Wealden's tiniest tetrapods - indeed, some of the smallest fossil tetrapods of all - facing off in leaf litter. Recently named Wesserpeton evansae, these are albanerpetontids, very small amphibians which only died out a few million years ago. The 35 mm snout-vent length of these animals did nothing to temper their ferocity, and numerous jaws of Wesserpeton have healed fractures and breaks from intraspecific tussles. The animals in this picture are speaking the aggressive body language of modern salamanders as a prelude to their conflict. Two sauropods hang around in the background because, hey, it's called the Age of Dinosaurs for a reason. Some people have suggested this image borders on the trippy and surreal. Stay off the shrooms, kids. 

Rebbachisaurids and chums

Lower Cretaceous rebbachisaurids and giant sauropod 'Angloposeidon' look for water in this desiccating Wealden lake.

I do like rebbachisaurids, that group of sauropods who didn't get the memo about long necks. They're only represented by scrappy remains in the Wealden (a scapula) which is enough to tell us they were there, but not substantial enough to carry a name. Here, a few individuals are digging around a rapidly drying lake-bed to find a substantial source of water: digging elephants were the inspiration for this scene. In the background, probable brachiosaurid 'Angloposeidon' struts its stuff. It's meant to be walking particularly tall - I like the idea that fossil animals would carry themselves in different, characteristic ways, just as modern animals do. A pink gnathosaurine pterosaur has snuck into the foreground, just because. 

A lesser-seen Wealden scene: the Hastings Beds palaeobiota

Finally for now, here's one more new painting. This is a reconstruction of a swollen river representing part of the Hastings Beds, the oldest deposits of the Wealden, complete with local reptile fauna. The animals shown here are really poorly known: titanosaur 'Pelorosaurus' becklesii (bits of forelimb), possible carcharodontosaurian Becklespinax altispinax (three dorsal vertebrae), eucryptodiran turtle Hylaeochelys belli (a shell), and the possible azhdarchoid previously known as 'Palaeornis cliftii' (humerus). So yes, take the 'restorations' of these animals with an evaporite mine of salt: they're really just better known, fairly 'generic' representatives of groups represented by these Wealden taxa, air-dropped into a Wealden setting. Becklespinax is obviously modelled closely on Concavenator, as they seem to be pretty closely related and have a similar taste in dorsal ornamentation. I gave Becklespinax a more vertical anterior sail margin however, as indicated by the fossil. There's an article waiting to be written on palaeoart like this - should we even bother 'reconstructing' poorly known scenes and species? I clearly think we should, but we'll have to discuss the reasons why another time. 

I'm just now realising that there's a lot of confrontation in these images. Come back soon for a more placid, relaxed set of pictures in part 2...

Episode 3: Bernissartids, the button-toothed Crocodyliformes

3/3 - this, ladies and gentlemen, is the end. At least, until the inevitable prequels where I'll ignore the canon of the expanded universe and do my best to tarnish everything you liked about the original trilogy.

Here we are then, the last instalment of the Wealden Crocodyliformes Trilogy. Following the posts on atoposaurids and goniopholidids, today we're going out with a bang by covering a newly described Wealden crocodyliform unleashed on the world this morning. The study was written up by my University of Portsmouth chums and colleagues Steve Sweetman, Ulysse Pedreira-Segade and Steven Vidovic (Sweetman et al. 2014), and Steve V. has covered some aspects of his involvement at his blog. The paper is open-access so, for the full skinny on the discovery, you should head here.

This most recently identified Wealden crocodyliform is among the most sophisticated and unusual of all Wealden crocs. Named Koumpiodontosuchus aprosdokitii, it is known from a well-preserved skull which was recovered in circumstances owing much to chance and good fortune (Sweetman et al. 2014). This animal is currently only known for certain from the Wessex Formation of the Isle of Wight, specifically from fossil-rich cliffs next to the seaside village of Yaverland, and the only known skull of it is broken in half. The posterior half was discovered in March 2011 by holidaying fossil hunters, who took it to the local dinosaur museum (Dinosaur Isle, of Sandown) to have it identified. Another family, on a fossil-hunting holiday three months later, then found the front half of the skull. They took this to the same museum where, by chance, the same museum staff who’d handled the first piece were on hand. It was realised that each piece belonged to the same specimen, and the first half was rapidly brought back to the museum to check the degree of articulation. Remarkably, the join between the broken pieces was near perfect – clearly neither chunk had been exposed to weathering effects very long before being discovered – and the entire skull could be seen. Each piece was then donated to the museum to allow its study. Given the chain of events and people involved in the discovery of Koumpiodontosuchus, it’s easy to imagine how only single halves of the skull might be known to science, or even neither. This is clearly yet another story which stresses the importance of amateur fossil hunters to Wealden fossil discoveries, and the benefits of responsible collecting.

Holotype skull and mandible of the button-toothed crocodyliform, Koumpiodontosuchus aprosdokitii. From Sweetman et al. 2014.

Button-toothed crocodiles in context

Koumpiodontosuchus is a member of Bernissartidae, a group named by Sweetman et al. (2014) which only contains two species: Koumpiodontosuchus and Bernissartia fagesii. The latter is a famous, small Jurassic and Cretaceous crocodyliform known from France, Denmark, Spain, Portugal and particularly Belgium, where a spectacular complete skeleton has been unearthed. Indeterminate species of Bernissartia also seem to occur in the Ashdown Formation of Hastings (Salisbury and Naish 2011), but this identification may eventually warrant reappraisal now that Koumpiodontosuchus has been discovered. Bernissartid remains are not new, some of the first material of these animals being documented in the 1850s and Bernissartia itself being named from Belgian fossils in the 1880s. Isolated teeth, likely referable to Koumpiodontosuchus, have been found in Wealden deposits since at least the 1970s (Buffetaut and Ford 1979), so were clearly present across the entire geographic and stratigraphic range of the Wealden Supergroup.

Bernissartia has long been a bit of an oddball among Crocodyliformes, possessing some unusual anatomy and being of uncertain placement in crocodyliform systematics. The discovery of Koumpiodontosuchus provided a bit of light on this front, suggesting that Bernissartia was part of a group containing at least one other similar species, and that they occupy an evolutionary place between atoposaurids and the goniopholidid + Eusuchia radiation. This position isn’t too surprising, as there are a number of features in bernissartids which link them to Eusuchia – see below. Bernissartidae is primarily defined by dental characteristics, with the most obvious one also being the namesake of Koumpiodontosuchus: “button-toothed crocodile” (if anyone wants a common name for these Crocodyliformes, this is the one to use). The posterior teeth of bernissartids are rather globose – wide, short and blunt – and distinctive compared to the dentitions of most other Crocodyliformes. It’s these teeth which, even in isolation, betrayed the presence of bernissartids in the Wealden well before the more substantial Koumpiodontosuchus fossil was discovered. Their other teeth are quite different to this, however. The mid-region dentition is rather conical in shape; ‘pseudocanines’ erupt about 25 % of the jaw length from the jaw tip, and conical teeth emerge procumbently from the jaw tips themselves. Koumpiodontosuchus has two large pseudocanines on its lower jaw, which erupt so close to each other that they share a single, enlarged tooth socket. Bernissartia, by contrast, only possesses one.

The new Wealden bernissartid Koumpiodontosuchus aprosdokitii foraging for molluscs. It's eating a mud snail, Viviparus cariniferus, while tiny (6 mm long) physid gastropods Prophysa crawl over pond scum in the lower left of the image. Dragonflies provide scale, while unnamed tetanurans (based on findings of Benson et al. 2009) prowl around the background. An earlier version of this reconstruction was featured in Sweetman et al. (2014). Prints of this image are available here.

Bernissartids packed this sophisticated dentition into relatively tiny jaws: these were not big crocodyliforms. Indeed, with body lengths of approximately 600 mm, bernissartids were probably the smallest crocodyliform species in the entire Wealden succession. Like goniopholidids, bernissartids bore osteoderm shields on their backs and bellies, but the dorsal series was rather more complex than those of other Wealden crocodyliforms. Rather than possessing two rows of interlocking osteoderms as we saw in goniopholidids and atoposaurids, bernissartids possess four rows of osteoderms along their backs. These comprise two sets of rectangular, double-keeled scutes along the midline, and laterally bordering square osteoderms with single keels (Salisbury and Frey 2001). None of these interlocked, and – based on what we’ve discussed for other Wealden Crocodyliformes – it’s worth considering what impact this had on bernissartid locomotion. Rather than supporting their trunks with scutes, it seems that bernissartids developed procoelus trunk vertebrae (that is, vertebrae with centra extending into the corpus of the vertebra behind) to support their bodies when walking (Salisbury and Frey 2001). This feature, along with their relatively complex osteoderms, is shared with eusuchians and are some of the reasons why these animals have classically been allied to these Crocodyliformes. Of further interest here is the biconvex nature of the first bernissartid tail vertebra – this has further implications for their locomotion, which we’ll get to below.

The bit on palaeoecology

Ecologically, it seems that bernissartids had a preference for hard shelled prey. Their blunt posterior dentition has been labelled as ‘tribodont’ – literally meaning ‘crushing teeth’ – and, like slamming a couple of anvils together, are ideally shaped to crunch hard shells. Some confirmation of this idea is seen in the wear facets often seen on tribodont bernissartid teeth. Classically, their prey was largely considered to comprise molluscs such as the freshwater snails and clams populating Wealden streams and lakes (Buffetaut and Ford 1979). Recently, a broader diet has been postulated for bernissartids however, the logic being that hard shells are hardly restricted to molluscs even in freshwater settings (Sweetman et al. 2014). Insects and crayfish probably formed as much of their diet as molluscs, all of which were likely procured or extracted from soft-substrates with the procumbent anterior teeth. We should not forget the savage-looking pseudocanines of these animals however: these would be of little use against hard prey items, but may have allowed for spearing relatively soft-animals. Perhaps bernissartids are best viewed as rather opportunistic feeders, primarily taking hard-shelled prey but not turning their noses to other types of food when the opportunity arose.

If gastropods like this Wealden mud snail, Viviparus cariniferus, had nightmares, they contained bernissartids. 

Where was most of this prey caught? There is evidence that bernissartids were equally at home in water and on land. Their biconvex first tail vertebra suggests their tails were capable of considerable movement for providing burst propulsion through water and, unlike most other Wealden Crocodyliformes, their lack of interlocking osteoderms facilitated lateral trunk motion (Salisbury and Frey 2001). While compromising overall speed, this may have permitted greater amounts of manoeuvrability – ideal for pursing nimble, if relatively slow, aquatic arthropods. We’ve already mentioned that the reinforced trunk vertebrae of bernissartids would provide ample reinforcement for terrestrial locomotion, and their small size is relevant here as well. Like the small-bodied atoposaurids, and unlike the big goniopholidids, bernissartids had relatively small amounts of weight to lug around on land and could likely sustain long periods of terrestrial locomotion without tiring. It’s possible, therefore, that they found much of their prey on land as well as in water, perhaps enjoying the beetles, cockroaches and other tough-shelled terrestrial insects known to occur in Wealden deposits.

It’s worth pointing out that bernissartids may not be the only Wealden Crocodyliformes adapted for hard-shelled prey. The poorly known, 1.5 m long Wealden eusuchian Hylaeochampsa vectiana also has large posterior teeth ideal for smashing shelled prey (Clark and Norell 1992), although the dentitions of other hylaeochampsids are complex and it’s possible Hylaeochampsa had a very varied diet. As discussed for other Wealden Crocodyliformes, it’s likely that the size difference between the bernissartids and Hylaeochampsa would prevent too much overlap in prey preference: the latter may have been capable of eating large molluscs or even small armoured vertebrates, which were probably unavailable to bernissartids. There's lots more we could say here, but I'd best not - maybe Hylaeochampsa will warrant dedicated discussion at a later date.

The end

And I guess that's where we'll leave the Wealden Crocodyliformes for now. As alluded to above, there are other crocodyliform species and groups we could discuss, but they're generally less well known than the taxa we've covered across these posts and it would be difficult to discuss them in comparative depth. I hope you've enjoyed this series of themed posts and, if artwork of ancient Wealden animals is your thing, come back soon for a big announcement about an event related to just that.

References

• Benson, R. B., Brusatte, S. L., Hutt, S., & Naish, D. (2009). A new large basal tetanuran (Dinosauria: Theropoda) from the Wessex Formation (Barremian) of the Isle of Wight, England. Journal of vertebrate Paleontology, 29(2), 612-615.
• Buffetaut, E., & Ford, R. L. E. (1979). The crocodilian Bernissartia in the Wealden of the Isle of Wight. Palaeontology, 22(4), 905-912.
• Clark, J. M., & Norell, M. (1992). The Early Cretaceous crocodylomorph Hylaeochampsa vectiana from the wealden of the Isle of Wight. American Museum novitates; no. 3032.
• Salisbury, S. W. & Naish, D. (2011). Crocodilians. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 305-369.
• Salisbury, S. W. & Frey, E. 2000. A biomechanical transformation model for the evolution of semi-spheroidal articulations between adjoining vertebral bodies in crocodilians. In Grigg, G. C., Seebacher, F. & Franklin, C. E. (eds) Crocodilian Biology and Evolution. Surry Beatty & Sons (Chipping Norton, Aus.), pp. 85-134.
• Sweetman, S.C., Pedreira-Segade, U., & Vidovic, S. (2014) A new bernissartiid crocodyliform from the Lower Cretaceous Wessex Formation (Wealden Group, Barremian) of the Isle of Wight, southern England. Acta Palaeontologica Polonica (in press)

Episode 2: The Wealden River Masters, goniopholidid Crocodyliformes

Insert your own whoops, hollers,cheers, or discharging firearms here. 

Welcome to Episode 2 of the snappily-titled Wealden Crocodyliformes Trilogy!* We'll waste no time with introduction - read this if you haven't already - and dive straight into our second group, the goniopholidids. Much of the information herein is derived from Salisbury and Naish (2011) so, if in doubt, consult this tome for further details.

*Snappily titled? And it's about crocodiles...? Man, I'm so wasted on you guys.

Without question, the Wealden waterways were lorded over by a group of Crocodyliformes known as Goniopholididae. The namesake of this group, Goniopholis, is one of the more familiar Mesozoic crocodyliforms after famous taxa like Sarcosuchus and Deinosuchus, and is well known as a relatively ‘conventional’ crocodyliform compared to some of the other weirdo crocs doing the rounds in the Mesozoic. Goniopholidids are found throughout Jurassic and Cretaceous rocks in the Northern Hemisphere and are part of several famous fossil faunas, including being the best known crocodyliforms of the Wealden fauna. Miscellaneous goniopholidid teeth and scutes occur throughout the Wealden, and their existence has been known for a long time. Teeth ultimately attributed to indeterminate goniopholidids were found in Sussex during the 1820s by Gideon Mantell as part of the same collections of crocodile’ material which was later found to contain unappreciated early records of Wealden baryonychines.

Despite this long history, work on Wealden goniopoholids is still developing (Salisbury and Naish 2011). At one time, most Wealden goniopholid taxa were considered members of Goniopholis proper, the famous Owen-named genus of great historic significance. As with many 'classic' genera, Goniopholis is now appreciated to be a bit of a taxonomic mess and claims of 19 species are being scrutinised (e.g. Salisbury and Naish 2011; Andrade et al. 2012). Recent reviews have suggested that the Wealden goniopholidid assemblage contains a sole Goniopholis species from the Weald Sub-basin and two other genera from the Wessex Sub-basin, all known from good skull material and, in the latter instances, a series of articulated postcranial remains (Salisbury and Naish 2011). These named species include Goniopholis willetti from the Grinstead Clay Formation, Sussex; Anteophthalmosuchus hooleyi (below), from the Wessex and Vectis Formations of the Isle of Wight, and Vectisuchus leptognathus, also of the Wessex Formation (why Vectisuchus when it’s not found in the Vectis Formation? ‘Vectis’ is the Roman word for the Isle of Wight, so ‘Vectis’ frequently pops up in animal names from this part of the world). The latter was almost known from a complete skeleton, but a cliff fall during its collection rendered much of the hindlimb, pelvis and tail inaccessible. In spite of this incident, it’s still fair to say that this group has one of the better records among Wealden reptiles, and it might get even better. Fragmentary goniopholidid jaw fossils hint at further, unnamed species, but they are currently too poorly represented to warrant naming.

The Wealden goniopholidid Anteophthalmosuchus hooleyi takes advantage of a flooding river to hunt two stranded Hypsilophodon foxii. The big one is Using the Ballet to escape. Prints of this image are available here.

Goniopholidids vs. modern crocodilians, round 1: anatomy

What kind of Crocodyliformes were goniopholidids? Because these animals appear to resemble modern crocodiles in size, shape and probably lifestyle moreso than any other well-known Mesozoic Crocodyliformes, they are often reconstructed as ancient copies of large modern species like Nile or saltwater crocodiles (e.g. Karl et al. 2006 - see reconstruction here). This isn’t really the case, however: goniopholidids may look a little similar to modern crocodilians at first glance, but much of their anatomy is unconventional and their possible habits were likely rather different. If we compare these aspects directly, their differences will soon become apparent.

We’ll start with size. Here, it must be said, goniopholids are undoubtedly pretty similar to modern crocodilians. The largest Wealden goniopholidids – Anteophthalmosuchus and G. willetti - were large animals each attaining at least 3.5 m long. This is a pretty comparable size for many modern crocodiles, and may even seem a little on the small side compared to the 5 m+ lengths attained by some extant crocodilians. Don’t be fooled into thinking this makes Wealden goniopholids diminutive creatures, however: a 3.5 m long crocodyliform would somewhere around 200 kg in weight and stretch longer than your 3-seater sofa. These were undoubtedly big, bulky animals. Vectisuchus, by contrast, was a much smaller species, only attaining 1.2 m in length.

In fine anatomy, we start to see obvious differences between the ancient goniopholids and modern crocodilians. Goniopholidid backs were covered with two rows of rectangular osteoderms with interlocking pegs at their distal margins – they are much like the atoposaurids we met last time in this respect (Salisbury and Frey 2000). These are largely devoid of ornamentation with only slight keels along their dorsal surfaces. This configuration is rather different to the more complex and ornate osteoderm arrangements seen in modern crocodyliforms, and goniopholidid osteoderm shields would probably seem rather simple and inelegant by contrast. Also unlike modern crocs, goniopholid osteoderms do not extend far up the neck, perhaps because doing so would impair neck mobility (see below), and further osteoderms were found along their bellies. These were formed of hexagonal plates rather than long, rectangular ones however. Another key distinction between goniopholidids and modern crocodilians is found in their forelimbs. Most goniopholids have arms which are at least as long as their legs and many species - including Anteophthalmosuchus and Vectisuchus – have forelimbs which surpass the length of the hindlimb. This increased length is provided by relatively elongate humeri and wrist bones, and would give goniopholidids taller statures than those of all modern crocs, which are always shorter up front than behind. If we extending this comparison further, we’ll see that goniopholidid forelimb length is almost unique among all Crocdyliformes, being longer than virtually all of their relatives.

Like many modern crocodilians, goniopholidids possess the well-built, powerful skulls of formidable predators. There is also overlap in general skull shape with modern crocs too, with G. willetti and Vectisuchus having rather long, slender snouts which are narrower than the posterior regions of their jaws. By contrast, the skull of Anteophthalmosuchus belongs to a real bruiser; its jaws only gently converge from the enormous posterior region to form a chunky, roughly triangular skull with a rounded muzzle. Both skull types are equipped with goodly-sized, slightly recurved conical teeth which would not look out of place on modern crocodilians. Again however, there are differences in detailed anatomy. Of particular interest is the orbits of Anteophthalmosuchus, which only permitted forward vision rather than anterolateral as is usual for Crocodyliformes (its name, roughly meaning ‘forward-eye-crocodile’, reflects this - see Salisbury and Naish 2011). A similar condition is also seen in Vectisuchus, but it is not quite as well developed and seems to have arisen independently. Goniopholid skulls are also rather flatter than those of modern crocs, and have distinctly over-biting upper jaws. An unusual hollow in the cheek region, known as the maxillary depression, was also present, apparently representing an unusually large pressure-sensitive region of the goniopholidid face (Andrade 2009).

A house-proud Goniopholis willetti stands at the entrance to his burrow. Note his long arms, narrow jaws, and lack of a doormat.

Goniopholidids vs. modern crocodilians, round 2: habits

It may be expected that these similarities and differences between modern crocodilians and goniopholididis may translate to overlapping, but also slightly different lifestyles. Happily, because the anatomy of Wealden goniopholidids is well-documented, we can make some informed speculation as to how these animals may have lived and, indeed, this seems to be the case. The size and robust skeletons of G. willetti and Anteophthalmosuchus suggests that the ecological bucks of Wealden waterways stopped with them: occasional visits from spinosaurids aside, they were the largest predators in Wealden lakes and rivers and clearly well suited for tackling large prey items. We might imagine each as the apex predators of their respective waterways, taking small or medium-sized terrestrial animals, large fish and other aquatic reptiles as prey. This gives these animals a role much like those filled by several species of large crocodilians today. Smaller Vectisuchus, by contrast, probably ranked it in the mid-league of ancient Wealden ecosystems, probably capable of holding its own against most aquatic Wealden species but wanting to be wary of its larger cousins. Applying trends of snout shape and prey preference of modern crocodiles to Wealden goniopholidids suggests they likely differed in general prey preference: slender-snouted Vectisuchus and G. willietti probably took relatively smaller prey than the massively-jawed Anteophthalmosuchus. Through overall body size and jaw shape, these animals probably avoided stepping on each other’s ecological toes – at least Anteophthalmosuchus and Vectisuchus were contemporaries which probably practised niche partitioning (Salisbury and Naish 2011).

We might expect goniopholidids to exploit their large size in a similar way to modern crocodilians. Large modern crocodiles often focus their predation efforts to certain times of year when environmental conditions are favourable, such as times when rivers and lakes are in flood, when prey is particularly abundant, or at least the climate is more forgiving. Given how extreme the Wealden climate was - summer temperatures in some parts of the Wealden reached 36–40°C and experienced annual droughts (Sweetman and Insole 2010) – goniopholidids may have used similar strategies. As with big modern crocodilians, their large bodies hold ample reserves to wait out leaner or stressful times, and it’s possible that some goniopholidids waited out the long, hot Wealden summer in cooling pools or burrows (see image, above), while smaller crocs had fewer resources to fall back on and continued to exert themselves throughout hard times.

Beyond these generally favourable comparisons however, many aspects of goniopholidid anatomy hint at different habits to modern crocodilians. For instance, the development of goniopholidid maxillary depressions likely represent enlargements of sensory organs present in modern crocodilians used to detect prey at the water/air interface (Andrade 2009). All else being equal, does this indicate that goniopholids were more routinely grabbing prey at the water surface rather than diving for food or living generalist lifestyles? In other instances, it’s not clear what significance goniopholidid anatomical quirks may have. It’s difficult not to wonder why some Wealden goniopholidids possess entirely forward-facing eyes, for instance, and if this was related to feeding. Ordinarily, increased amounts of forward vision are associated with development of binocular vision and heightened abilities to judge distances. Might that mean predation techniques were unusual in some goniopholidids, involving chases, or carefully judged lunges and strikes at prey?

The preferred habitats and locomotory methods of goniopholidids are also worth pondering. There is some evidence that larger Wealden goniopholidids were mostly confined to a semi-aquatic existence, as their interlocking osteoderms likely strengthened their backs and improved terrestrial competency (as it does for atoposaurids and several other type of ancient crocodyliform), but their sheer weight likely impeded terrestrial locomotion over sustained periods (Salisbury and Frey 2000). The same is true of large modern crocodiles: here, reinforced vertebral joints perform a similar job to osteoderm bracing but still fail to facilitate effective, fast terrestrial locomotion for long periods. Larger crocodilians therefore spend much of their time in water, and certainly find most of their food there. If so, this makes the atypically long forelimbs of goniopholidids all the more interesting. Often, development of relatively equate limb lengths in quadrupeds is considered a sign of good terrestrial proficiency, betraying a well-balanced animal with effective carriage and equal gait efficiency in both limb sets. But how can this apply to large, heavy goniopholidids if they weren’t walking very much? Doubtless, an increased forelimb stride length was useful on occasions when large goniopholidids did leave the water, but why develop these features if much of their lives were spent in deep water? Did these animals ‘walk’ along river beds more than other Crocodyliformes? Was this trait even important for big adults? Perhaps smaller or juvenile goniopholidids took advantage of long forelimbs before they outgrew real terrestrial proficiency, spending more time on land before becoming more thoroughly aquatic at larger sizes. We could speculate all night about the intriguing possibilities here: Crocodyliformes are sophisticated creatures which do a lot more than eat, sleep and wander about: they also dig burrows, construct nests, climb onto trees and rocks, and are very sociable. Could their long forelimbs be related to these behaviours? Vertical size is seemingly more intimidating to modern Crocodyliformes than girth (Farlow and Dodson 1975) - might long arms and a tall stature have incurred social significance for goniopholidids? It’s not inconceivable that the long arms of goniopholidids were influenced by these activities rather than just locomotion, and I suspect an investigation into the evolution and functionality of their forelimbs would yield some very interesting results.

The outcome

In sum, then, it seems that we need to be cautious when thinking of goniopholidids as 'conventional' Crocodyliformes or simply forebears of modern crocodiles. Many aspects of their anatomy are not only different from those of modern crocodiles, but actually downright odd, and likely impacted on their habits and lifestyles significantly. Palaeoartists - bear all this in mind the next time you set out to draw your goniopholidids skulking in the background of your dinosaur artwork.

For the concluding post in the Wealden Crocodyliformes Trilogy, we're going out with a bang and an exciting new discovery - and it's not very far off now. Stay tuned!

References

• Andrade, M. B. (2009). Solving a century-old mystery: the structure and function of the maxillary depressions of Goniopholis (Crocodylomorpha, Neosuchia). In Journal of Vertebrate Paleontology (Vol. 29, pp. 54A-55A). 
• Andrade, M. B. de, Edmonds, R., Benton, M. J., & Schouten, R. (2011). A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus. Zoological Journal of the Linnean Society, 163(s1), S66-S108.
• Farlow, J. O., & Dodson, P. (1975). The behavioral significance of frill and horn morphology in ceratopsian dinosaurs. Evolution, 353-361.
• Karl, H. V., Gröning, E., Brauckmann, C., Schwarz, D, & Knötschke, N. (2006). The Late Jurassic crocodiles of the Langenberg near Oker, Lower Saxony (Germany), and description of related materials (with remarks on the history of quarrying the “Langenberg Limestone” and “Obernkirchen Sandstone”). Clausthaler Geowissenschaften, 5, 59-77.
• Salisbury, S. W. & Frey, E. 2000. A biomechanical transformation model for the evolution of semi-spheroidal articulations between adjoining vertebral bodies in crocodilians. In Grigg, G. C., Seebacher, F. & Franklin, C. E. (eds) Crocodilian Biology and Evolution. Surry Beatty & Sons (Chipping Norton, Aus.), pp. 85-134.
• Salisbury, S. W. & Naish, D. (2011). Crocodilians. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 305-369.
• Sweetman, S. C., & Insole, A. N. (2010). The plant debris beds of the Early Cretaceous (Barremian) Wessex Formation of the Isle of Wight, southern England: their genesis and palaeontological significance. Palaeogeography, Palaeoclimatology, Palaeoecology, 292(3), 409-424.