#teleosauroidea

[arminreindl]

Croc Paleontology Recap January 2025

The year has just begun and already we got a bunch of pretty neat new studies on fossil pseudosuchians so I'll just briefly go over them and unless I forget or end up procrastinating/getting too busy I'll hopefully be able to keep this going throughout the rest of the year.

Just to give you a brief overview, the highlights of this month include: salt glands in gavialoids, crocodilian predation on azhdarchids, diversifications and extinctions in thalattosuchians and a new species of aetoaur from India. Lets begin.

Evidence for salt glands in gavialoids

Starting off with something relating to the dispersal of gavialoids, we got "Evaluation of the endocranial anatomy of the early Paleogene north African gavialoid crocodylian Argochampsa krebsi and evolutionary implications for adaptation to salinity tolerance in marine crocodyliforms" by Pliggersdorfer, Burke and Mannion.

The title already gives a lot away, but the point was that Argochampsa, from the early Paleocene Ouled Abdoun Basin in Morocco, was examined for evidence of salt tolerance. Why? Because the dispersal of gavialoids remains weird. Both modern forms aren't especially keen on saltwater and are only known to consistently occur in freshwater (tho we have a recent example of an indian gharial caught in a fishing net off the coast of India), yet we have plenty of extinct gavialoids that either indicate that the group must have crossed oceans (see any "gryposuchine") or straight up lived at sea (also see some "gryposuchines").

Now, one such example might also be Argochampsa. Both because the Ouled Abdoul Basin famously preserves coastal deposits and because, at least following some phylogenies, Argochampsa might be closely related to the gharials of South America and today (others say its not even a gavialoid but lets ignore that for now). So all things considered one might expect marine habits from Argochampsa, yet so far no such adaptations could be identified. Well, Pliggersdorfer and co. analyzed a thus-far undescribed skull and actually managed to find something. Small depressions on the inside of the skull are suspiciously similar to ones seen in the extinct, fully marine metriorhynchoids, depressions that in the latter have been interpreted as having been left by salt glands. There is also some further evidence through the morphology of the inner ear.

This conclusion further extends to a handfull of other taxa, including the dyrosaurid Rhabdognathus and the recently named gavialoid Sutekhsuchus, and lends itself to the hypothesis that salt glands may have been ancestral to gavialoids, something I personally find unsurprising given their proximity to crocodyloids and their dispersal across the world (really if anything alligatoroids seem like the odd one out).

Fun fact, yours truly is featured in the paper in the form of two silhouettes.

Left: Argochampsa, illustrated by Seismic Shrimp/JW Right: Piscogavialis, perhaps the most famous marine gavialoid, illustrated by Joschua Knüppe

The brain of Paralligator

Second on our neat little list, the neuroanatomy of Paralligator, studied through CT scans and 3D modeling and published on in "Neurocranial anatomy of Paralligator (Neosuchia: Paralligatoridae) from the Upper Cretaceous of Mongolia". Given that I am not great with brain things, I'll keep this one short.

Now for those unfamiliar, Paralligator is part of a somewhat strange clade known as the Paralligatoridae, which contrary to their name are nowhere near real alligators (tho some do look deceptively similar). Instead, they are much more basal members of Eusuchia.

Measurements of the olfactory bulbs, responsible for the sense of smell, indicate that in Paralligator this sense was similarily developed to allodaposuchids and crocodilians, as is the inner ear who's anatomy suggests a semi-aquatic lifestyle. Paralligator does however differ in possessing a mesothemoid, a bony septum in the olfactory region that is also seen in dyrosaurids, baurusuchids and dinosaurs, but not modern crocodiles.

Borealosuchus remains from Colorado

Tho seemingly unexciting, this study, "First record of Borealosuchus sternbergii from thelower Paleocene Denver Formation (lower Danian),Colorado (Denver Basin)" actually helps us fill a neat little gap in our previous knowledge on croc survival across the Cretaceous-Paleocene extinction.

Simply put, though America's croc record across the KPG is rather remarkable, showing both many survivors and some incredible diversification after the impact, Colorado is kind of a blind spot, despite its potential importance. Perhaps one of the best examples of a survivor concerns the genus Borealosuchus, which is both geographically and stratigraphically widespread. To put things into perspective, this genus occured as far north as Canada and as far south as Texas, first appearing in the Late Cretaceous and dying out in the Eocene.

This paper now described several skulls from the Corall Bluff's locality of the Denver Formation, earliest Paleocene, that can be attributed to Borealosuchus sternbergii, definitively extending its range beyond KPG (granted, there are tentatively referred Paleocene occurences elsewhere), making it one of the largest suvivors of the mass extinction, with adults growing up to 2.3 meter in length. The specimens from Colorado are smaller, in the 1.5 to 1.7 meter range, but they are also regarded as immature individuals and are therefore also regarded as usefull in illustrating how the animals changed as they grew into adulthood.

This paper is especially well timed for those that follow @knuppitalism-with-ue 's Formation Stream series. As you might know, Corall Bluffs is to be drawn barely a week from now and this is a fantastic addition.

Left: Borealosuchus drawn by Atak_Draws Right: Distribution of Borealosuchus by Lessner, Petermann and Lyson 2025

Growth of a peirosaur

Our next paper for discussion is "Life history and growth dynamics of a peirosaurid crocodylomorph (Mesoeucrocodylia; Notosuchia) from the Late Cretaceous of Argentina inferred from its bone histology" by Tamara G. Navarro and colleagues. This study conducted the first histology of peirosaurid limb bones, specifically of an indetermined taxon clading together with Uberabasuchus.

As a brief refresher, peirosaurids are a branch of medium to large sized Notosuchians that I personally think can be aptly described as appearing somewhat like scaly dogs or pigs with often robust, wedge-shaped heads and heavily armored bodies.

The results show that the animal had reached sexual maturity, yet was not yet fully grown. What's also noted is the exact growth dynamics of this animal. This is to say, the studied peirosaurid had overall slow growth with cycles of no growth whatsoever and two periods of increased growth, tho once put against other notosuchians the study deems the growth rates to be better described as "moderate". Pepesuchus meanwhile, belonging to the closely related itasuchids, was a fast grower. Extending things beyond their shared clade shows a virtual mish-mash of dynamics, with Araripesuchus buitreraensis displaying slow growth rates (yet Araripesuchus wegeneri having faster rates than the peirosaur), Iberosuchus showing slow rates, and Notosuchus displaying high growth rates (hell, theres even variation between individuals). A final point concerns the age of the individual, which is....contradictory. Based on the limb bones, the study estimates that the animal was at the very least 15 years old, but previous study of the osteoderms has yielded an estimated age of 18 years old. Ultimately, further study is needed, but it does clearly show how the histology of different parts of the skeleton varies.

Shown below, Uberabasuchus terrificus by Scott Reid

Predation on pterosaurs

Here's a fun one, "A juvenile pterosaur vertebra with putative crocodilian bite from the Campanian of Alberta, Canada", once again with a name that tells you very much what you're in for.

Brown and colleagues report on the discovery of a juvenile specimen of the azhdarchid Cryodrakon from the Dinosaur Park Formation of Alberta, Canada. The neck vertebra bears some conical bite marks, notably different from those of theropods, which generally have D-shaped or compressed tooth crosssections (sans spinosaurids, which aren't present). Champsosaurus is also ruled out due to its inferred feeding preferences, weak bite force and slender teeth. Mammals are potential candidates, but the team regards it as more likely that the trace maker was a crocodilian. Considering the fauna of the Dinosaur Park Formation, this would suggest the culprit was either Leidyosuchus, Albertochampsa or an animal described as "Stangerochampsa-like".

Now this is a very interesting, if not exactly unexpected interaction. On the one hand, having direct fossil evidence for this is a big deal, even if we don't know if the bite marks were left due to the pterosaur being actively hunted or if they were simply left when a lucky croc came across the carcass of an already deceased Cryodrakon. On the other hand, crocodiles and kin are notoriously opportunistic and broad in their diet, so one feeding on a pterosaur is something that seems like a no-brainer in principle, especially a relatively small individual with a wingspan of "only" 2 meters. This is further supported by the fact that crocodilian bite marks have also been reported from the Romanian pterosaur Eurazhdarcho.

Obvious difference in prey size and geography aside (and taxon names even within the chosen setting while we're on it), Prehistoric Planet really nailed the nailon the head with this one.

In the left corner, a juvenile Cryodrakton (art by Hank Sharpe). In the right corner, Leidyosuchus (art by Gunnar Bivens) LET THEM FIGHT (or scavenge)

Evolutionary trends and extinctions in Thalattosuchia

This one's a last minute entry, by which I mean this one got just published as I was about to wrap this whole thing up. "Analysing Thalattosuchia palaeobiodiversity through the prism of phylogenetic comparative methods" explores how the evolution and the extinctions of members of this group were shaped by both biotic and abiotic factors.

Given the shere breadth of this topic, a quick summary of thalattosuchia seems kinda in order. In short, thalattosuchians are a group of what are likely to be early crocodyliforms adapted to life at sea. They can be split into two groups, the teleosauroids, superficially gharial like animals that likely stuck to coastal waters, and the metriorhynchoids, open ocean animals with fluked tails, no body armor and paddle-like limbs. Both groups reached their greatest diversity in the late Jurassic, but managed to survive into the Cretaceous before disappearing entirely.

The study recaps that thalattosuchians first reach great diversity during the Toarcian, tho this is likely influenced by preservation bias thanks to Lagerstätten such as the Posidonia shale, and a later diversification takes place during the Bathonian. Regardless, the transition from the lower to middle Jurassic sees an increased trend in both thalattosuchian groups towards shorter snouts, which are associated with durophagy or hypercarnivory. This essentially gives rise to the teleosauroids of the Machimosaurinaei, which appear during the Bathonian and have blunt, robust teeth, as well as the metriorhynchoid Geosaurinae, which appeared at the same time and had ziphodont (serrated teeth). The reasons for this could be twofold. On the one hand, thalattosuchians were very abundant, so expanding into new nisches helped them coexist, with ecosystems preserving fish-eaters, hypercarnivores, durophages and more at the same time. More of an underlying factor could be a drop in ichthyosaur diversity, leaving plenty of open nisches for these crocs to fill.

Subsequently, during the transition from the Middle to Late Jurassic, there was another diversification event with both groups establishing new major clades, possibly associated with the warm temperatures of the Late Jurassic, before the diversity crashes with the onset of the Cretaceous. The authors note that this too might have been related to climate, with the Cretaceous survivors mostly being found in warmer waters.

Left: A Dakosaurus ambushing an ichthyosaur by Gabriel Ugueto Right: A large Machimosaurus rests on the beach as a sauropod approaches, art by Joschua Knüppe

and for the final study I wanna talk about

Kuttysuchus: A new Aetosaur from India

Now, by all accounts one might be surprised to see this just kinda thrown in at the end here rather than getting a dedicated post as I usually like to do with new forms. And truth be told, theres just not that much to say about "A New paratypothoracin aetosaur (Archosauria: Pseudosuchia) from the Upper Triassic Dharmaram Formation of India and its biostratigraphic implications".

Kuttysuchus is our first pseudosuchian to be described this year and to get things out of the way, its not super exciting in terms of material. Like some other recently named aetosaurs, Kuttysuchus is based entirely on a handfull of osteoderms. And there's nothing wrong with that, after all osteoderms are rather distinctive for these animals. It does however mean that the information we can get from them is a bit limited and thus makes it hard to really put together something engaging.

More interesting than the anatomy then is the range and its relationship to other aetosaurs. The fossils are known from the Dharmaram Formation of India, which you might recall is also home to the recently named Venkatasuchus. Both Venkatasuchus and Kuttysuchus are members of the Paratypothoracini, tho the former is significantly more derived and the latter more basal.

Fossil osteoderms of Kuttysuchus, all belonging to the central double row that stretches across the back.

I'll be entirely honest. This was a lot more work to type out than anticipated, but admittedly also fairly rewarding. Hopefully you dear reader found it equally interesting, and hey, congrats on making it to the end.

[arminreindl]

Two more croc-line archosaurs to dig into with this one and both are super fascinating.

Obviously the star of this piece is our friend in the foreground, Plesiosuchus manselii, the largest of the fully marine metriorhynchids (the skull of which being the first one shown in the image to the right, taken from Young et al. 2012)

More broadly speaking, metriorhynchids are a fascinating group of early croc-relatives that stand out for having returned completely back into the water, developing a fluked tail, tiny little reduced forelimps, flipper like hindlimbs and smooth skin entirely lacking in the osteoderm armour we so strongly associate with pseudosuchians.

Plesiosuchus specifically is the largest of them. Close to 7 meters in length, it's basically in the same range as Liopleurodon (although smaller than several of the pliosaurs we know from the Kimmeridge Clay like that very decorative skeleton). Plesiosuchus actually wasn't the only metriorhynchid from the formation either. We also know of the small Cricosaurus, the name-giving Metriorhynchus and most interestingly the large-bodied Dakosaurus and Torvoneustes, which were likely to reach lengths of 4.5 to 4.7 meters respectively. At least with metriorhynchids this diversity can be explained in the different animals having very different preferences. Torvoneustes is interpreted as a durophage, feeding on hard-shelled prey. Dakosaurus is characterized by slicing dentition, potentially able to cut apart large prey with tooth wear possibly suggesting sharks or suction feeding habits. Plesiosaurus meanwhile would be able to take on large prey simply due to its great size and large gape but may have been limited by the size of its own head.

The other marine "croc" in the image is Bathysuchus, which may be described as a cousin to the metriorhynchids.

Remember Machimosaurus from the Guimarota locality that Josch drew a few weeks back? Yeah its related to that one. A teleosauroid to be precise, which are the sister group to metriorhynchoids and appear much more croc-like in their appearance despite still being a long way off from crocodiles in the strict sense. Teleosauroids may appear like what you'd think of as basal to metriorhynchoids, but they simply did their own thing after diverging, thus retaining this more ancestral bauplan with more developed limbs, more croc-like skulls and maintaining their armour plating (keep that in mind).

Bathysuchus is kind of a weird one tho. Why? Because from what we can tell Bathysuchus seems to have tried to become more like its metriorhynchoid cousins. It's known from deeper waters, its osteoderms were much more reduced and if the closely related Aelodon is anything to go from then its quite likely that this guy also started reducing the size of its forelimbs. In short, adaptations to a more pelagic lifestyle that moves away from the coastal habitat other teleosauroids seem to have preferred. Who knows, maybe its possible that in some other timeline these guys might have become something akin to metriorhynchoids like what happened with mosasaurs.

Unlike metriorhynchoids, teleosauroids were way less common in the Kimmeridge Clay, hell, to my knowledge Bathysuchus is the only one of them, a pretender among the other pelagic pseudosuchians.

Result from the Kimmeridge Clay #paleostream!

We were only able to scratch the surface, but I think you get the idea ;)

[saritapaleo]

Patreon request for @/rome.and.stuff (Instagram handle) - Turnersuchus hingleyae

Described just last year, the Early Jurassic Turnersuchus is the most basal known member of the thalattosuchians: sea-dwelling crocodylomorphs. It existed before the two major thalattosuchian groups, the paddle-limbed Metriorhynchoidea and the long-snouted Teleosauroidea, split, and it may be the ancestor of both. While only parts of the skull are known, its shape suggests a narrow snout that was not as long as that of later thalattosuchians. Only one osteoderm is known, and it is oval with a slight keel down the middle, believed to have been attached to the tail. As humble as this small pseudosuchian was, it helped us fill in the major gaps of our understanding of thalattosuchia, and finally gave us an idea of their origin. Previously, all we knew was that the highly-derived metriorhynchoids and the more crocodile-like teleosauroids were related, lived alongside each other, and seemingly appeared in the Jurassic. While not as well known as the icthyosaurs, plesiosaurs, and mosasaurs, thalattosuchians were very successful sea crocodylomorphs until around the Early Cretaceous, and they owe their success to this little pseudosuchian who decided to return to the sea.

Found in the Charmouth Mudstone Formation of England, Turnersuchus would have lived alongside a variety of ammonites like Phricodoceras and Gemmellaroceras, as well as icthyosaurs like Leptonectes. It likely fed on the small fish that lived here. As it was not as highly adapted for ocean life as its descendants, it may have spent some time basking on land, where it could have come across early dimorphodontid pterosaurs and the armored Scelidosaurus.

[typhlonectes]

A new early diverging thalattosuchian (Crocodylomorpha) from the Early Jurassic (Pliensbachian) of Dorset, U.K. and implications for the origin and evolution of the group

Eric W. Wilberg,Pedro L. Godoy,Elizabeth F. Griffiths,Alan H. Turner &Roger B. J. Benson

ABSTRACT

Among archosaurs, thalattosuchian crocodylomorphs experienced the most extensive adaptations to the marine realm. Despite significant attention, the phylogenetic position of the group remains uncertain. Thalattosuchians are either the sister-group to Crocodyliformes, basal mesoeucrocodylians, or nest among longirostrine neosuchians. 

The earliest definite thalattosuchians are Toarcian, and already possess many synapomorphies of the group. All phylogenetic hypotheses imply a ghost lineage extending at least to the Sinemurian, and a lack of older or more plesiomorphic forms may contribute to the uncertain phylogenetic placement of the group. 

Here we describe a new species, Turnersuchus hingleyae, gen. et sp. nov., from the early Pliensbachian Belemnite Marl Member of the Charmouth Mudstone Formation (Dorset, U.K.). The specimen includes partially articulated cranial, mandibular, axial, and appendicular elements. It can be attributed to Thalattosuchia based on the following features: distinct fossa on the posterolateral corner of the squamosal; broad ventrolateral process of the otoccipital covering the dorsal surface of the quadrate; large supratemporal fenestrae lacking a flattened skull table; broadly exposed prootic; orbital process of quadrate lacking bony attachment with the braincase. This specimen represents the earliest thalattosuchian currently known from diagnostic material. 

Phylogenetic analyses of two published datasets recover Turnersuchus as the earliest diverging thalattosuchian, and sister to Teleosauroidea + Metriorhynchoidea. Bayesian tip-dating analyses suggest a Rhaetian or Sinemurian divergence of Thalattosuchia from other crocodylomorphs, depending on topology, with confidence intervals spanning from the Norian to the Pliensbachian. 

The new specimen extends the fossil record of Thalattosuchia, but the time-scaling analyses demonstrate that a significant ghost lineage remains.

Read the paper here:

https://www.tandfonline.com/doi/abs/10.1080/02724634.2022.2161909

[arminreindl]

Indosinosuchus the freshwater Teleosaur

Ignoring the fact that I skipped out on posting about my recent deepdives on Armadillosuchus and Odobenocetops (besides that one figure I made) on Tumblr, let me catch up by talking about Indosinosuchus.

Indosinosuchus (Indochinese Crocodile, after the Indochinese Tectonic Block) is a type of teleosaurid from the Late Jurassic Phu Kradung Formation of China. First of all the Phu Kradung is a fascinating formation with a long and somewhat convoluted dating history. For quite some time researchers argued whether or not its Late Jurassic (based on fauna) or Early Cretaceous (based on pollen). Recent papers seem to argue that both can be true, with the lower parts being Jurassic and the upper Cretaceous, which also accounts for a faunal turnover visible throughout the record. Related to Indosinosuchus, we have classically Jurassic teleosaurs in the lower levels and pholidosaurids in the upper.

artwork by Kmonvish Lawan and Sakka Weerataweemat

Whats far more interesting however is that the Phu Kradung Formation is continental in nature, preserving floodplanes dominated by lakes and rivers. This is quite the unusual habitat for a teleosaur, which are traditionally marine animals, living in the coastal or even open waters of the Tethys. In fact the only other known example of a freshwater teleosaur is an unnamed animal from China that sometimes clades closely to Indosinosuchus.

Indosinosuchus is known from a total of 10 fossil individuals, largely consistsing of skulls and mandibles with the occasional postcrania, which were all found in a single locality and have been determined to represent at least two different species: I. potamosiamensis (Thai River Indochinese Crocodile) and I. kalasinensis (Indochinese Crocodile from the Kala Province).

Now overall, Indosinosuchus had a pretty long and narrow snout compared to most modern crocodilians. However, among teleosaurs, this animal was actually among the more robust genera. Even more so when you consider that within teleosaurs, it falls within the teleosaurid family, not the overall more robust machimosaurids that would eventually become durophages and macropredators, cracking turtle shells.

Analysing the biomechanics of Indosinosuchus did show some cool things. The snout was pretty average at resisting stress and force, at least for teleosaurs, but the bite force was respectably high. The force that could be transferred to the tip of the snout in particular is among the highest in this group and the maximum force it could excert was the single highest among all teleosaurs.

The two species however begin to differ in how fast they could close its jaws. I. potamosiamensis follows what would be expected. A teleosaur with a massive bite but only closing its jaws slowly as in teleosaurs with similar mechanical advantage values. I. kalasinesis however could close its jaws surprisingly fast, something you'd otherwise expect from animals with a much lower bite force that needed to strike a lot more swiftly in order to catch prey.

While we don't quite know the precise ecology of the two species, scientists could deduce a few things. For one, obviously the difference in the jaw closing/opening speed is a good sign that they had somewhat different lifestyles, which could explain why two so similar species coexisted at the same time in the same place. Secondly, although its bite force was notably high, it does not appear like Indosinosuchus went after especially large prey. Sure, it could bite hard and had a relatively robust rostrum, but its teeth were not particularily specialised like in Machimosaurus. So it has been suggested that it was a generalist, not a specialist as shown in this figure by Johnson, Young and Brusatte.

Last but certainly not least is the way these animals died. It does seem rather odd that 10 specimens all seemingly died in the same place at the same time, with most preserving the skull and not much else. Well there may be a reason. One clue stems from the most complete non-skull we have, which is a remarkably well preserved spinal column preserved in articulation, but missing the skull. So the idea is that the carcass may have floated in rather still waters (like an oxbow lake) and the weight of the head would eventually separate the two pieces. The spine must have been burried rather quickly afterwards so that it wouldn't just fall apart. Well the hypothesis continues that this could have been during a drought, when water levels were gradually receeding. We know that modern crocodiles will amass in great numbers during droughts until bodies of waters are nothing but mud and crocs, so it makes sense here. The full drying out of their sanctuary would then contribute to preservation and we even have evidence that they spend a minimum of 8 weeks in the dry thanks to the feeding traces of scavenging beetle larvae.

There is also a second hypothesis that suggests that the bodies may have been washed on land during a flashflood, but I personally think the drought hypothesis is a lot more interesting. Plus I managed to get @knuppitalism-with-ue to draw something akin to the later, with an unfortunate group of Indosinosuchus being stuck in a dried up lakebed as a local mamenchisaur approaches in the search for water.

Obligatory link to the Wikipedia page post-overhaul: Indosinosuchus - Wikipedia and the before and after, wondering if I should do the same with Armadillosuchus and Odobenocetops or if I should just come back once I got the next project done (Doratodon)