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Seven Rays

June 13, 2017

Basically the idea I got for the main villains of Fylgia Twelve, a project I’ve been working on for a while. They’re basically reverse homunculi (FMA), since they embody virtues. They were created by Pazro, and follow his esoteric orders.

Diligence is the main antagonist by virtue of doing most of the work and facing the heroes most often. Resembles a stereotypical angel, except with a bird mask. Is very smart and has photokinesis relating to technology.

Humility is the de facto leader (to which even Pazro concedes frequently), by its sheer appreciation of everyone’s efforts. Is basically a seraphim, a mass of light and flame like a six-winged serpent.

Charity is the defector, finding Pazro’s endgame immoral, feeling that all deserve the best, including itself. Its photokinesis specializes in boosting allies, either healing them or defending them. Surprisingly anti-social and jaded, but always on the front-lines.

Chastity is probably the least likeable. Resembles an ecclesial man, and its photokinesis is specialized to restrain others physically. Is very self-righteous and borderline psychotic.

Kindness is a knight archetype, that traps its opponent in a single combat with itself, with the only way out being through death or by passing the surrounding “flames of love”. Through it, Kindness will encourage you to confess and overcome your faults, and repay every attack you deal to it in kind.

Temperance is blind, and carries scales, using them to balance things; the sound of the shifting balance helps guide it around. Very calm, and this calmness extends to its magic: instead of restraining you physically like some other Seven Rays, it will dim your physical sensations and desires.

Patience is the most idealistic of the bunch (which is saying something) and serves as their PR department. It avoids fighting whenever possible, but when it does get involved expect FTL, which allows it to fight unimpeded as well as prevent imminent disasters,

Aquamarine and Steven: The Hatred That Transcends The Doors of Tragedy

May 9, 2017

It was a very bad day for Steven. He was separated from his most true and exuberantly breasted love, the Amethyst.

Aquamarine, a devil ugly lepidopteran WHORE, had desired Steven for herself and kidnapped him! All the gems were left on earth, cursing the heavens for the theft and rape (in the stealing kind, not the sexy kind,,,, yet), for each wanted Steven for themselves. The king of the gods, Dyaus Pita, heard their wailing, and punished them by turning the scorned women into wild beasts, forever unable to love again. Even Amethyst, whose only crime was a broken heart, was forever a monster like mournful Medusa, and slain by Scarecrow with a spoon.

Now, Steven was lost in the vastness of the darkness of the emptiness of the blackness of the statelessness of star-less space. He was on the space saucer of the hubris-ful and arrogant Aquamarine, the gem who had captured him in name of the Diamonds, who wanted to wed him, but instead kept him for herself. Either fate was too monstrous for Steven, who loved Amethyst, who was now a monster, who was now killed by a kitchen utensil, by Scarecrow.

“Hahahahahahahahahahaha now you’re all mine my beloved Steven!” cackled Aquamarine conqueredly, crowing like a roster upon victimized serpents.

She could not help but be consumed by the lust of her beloved Steven, so she took out her wand and inserted it in her decadent and decaying uterus. Caterpillars and maggots gnawed on her flesh, now a massive gaping, pus-trimmed, necrotic hole sending directly to her womb, sending flashes of pleasure up her spine, who damaged her Broca area so each spasm was responded by bowel failure, ejecting globs of rotten black shit like a healthy pigeon (not the white guano of STARVED pigeons). She flew around the ship in ecstasy, throwing shitty everywhere in a choleric cholera. Her panties were completely submerged in brown, massive lumps falling slowly out of them as they were filtered by the tainted silk.

Her associates, the Topazes, sighed. They wanted Steven for themselves, but they had to serve the wretched fae, and so they took out a mop and began to washed the deck.

“Oh Steven, let us consummate our marriage!” said Mothman meanly, kissing the poor innocence boy with her putrid blackened gums.

Steven vomited in her mouth, his stomachal acids melting off her lips…but she LIKED IT EVEN MORE! She put a ring of gold with an amethyst gem on his finger, to remind him of the love he lost.

“Oh Steven, please your wife” moaned Aquaman moanfully, rubbing Steven’s underage nipples.

Steven sighed, he was obligated to copulate with the violator of his dreams.

But unknownst to them, Lars was hidden in the ship. He was very terrorfied, but also extremely jealous, because he wanted Steven for himself. So he walked to the Topaz, almost stepping on the poo.

“Hey big juicy lady” he said lyingly because he thought she looked like a lesbo megadyke, “Why do you take orders from that wimpy little fairy when you could take Steven as your husband?”

Topaz almost replied, but deep in her womanly heart she knew it to be true. So she threw the mop away and rushed towards Aukmarine. She was making out with Steven, so she was distracted and let her wand slip. Unfortunately, Topaz tripped on it and fell on them both!

“AAAAHHHHHHH YOU SHITTY ARNOLD IMPERSONATOR YOU ARE CRUSHING MY BONES!!!” roared Aquamarina annoyedly like that sun female dwarf from the Happily Ever After Hanna Barbera movie, but it was too late.

All her bones cracked, marrow infusing with her poo-tainted flesh. Most importantly, her skull was smashed against the pavement, and her false tear gem cracked, condemning her to a hell eternity of anal violation by symmetrodont concubines of yore. Unfortunately, Topaz weight also cracked Steven’s gem.

“Noooo, my love!” cried Topaz angstily, but it was of no use.

As Steven’s gem was destroyed, the flesh around his belly blackened and purpled with necrosis, ejecting pus like mayonnaise (geddit because its a gender and gems are non-binary). It spread all across Steven’s abdomen, dissecting his internal organs until they became jerky-like. His intestines were so compressed that all the poo inside him constipated, and constipated, and constipated until it was a massive bump on his belly. Steven moaned in agony, the pain was intense. With a single tear, he reached for the wand.

“For Amethyst, my one true love” he said, and committed seppuku.

In an instant, a pure black oily torrent erupted from his bowels like an ocean of darkness, washing away Topaz and Lars, dung filling their lungs and brains until they exploded. They joined Aquamarine in HELL, where they were most righteously tortured for ten billion years, boys and girls.

But what about our beloved Steven?

“Amethyst” was his last whisper, before his lungs deflated and fell into the poo.

Now Steven, Rose’s Son, was no more.

Then Samurai Jack and ASHI had SEX above his corpse.


Multituberculate Phylogeny

May 7, 2017


Pterosaurs and bats, hands and wings

May 2, 2017

Two diagrams by alphynix, showcasing the evolution of pterosaurs and bats. In both cases, the “missing link” is indeed missing.

As discussed previously, flying vertebrates most likely did not evolve from gliders. Not only does gliding not lead to powered flight (see this podcast for further examination on this in regards to birds), but we have evidence that bats evolved from fluttering, not gliding, ancestors (Padian 2011, Jepsen 1970), while pterosaurs evolved from ground-dwelling hopping ancestors (Witton 2015). The discussion on avian flight (as well as other dinosaurs like Yi) is still ongoing and far from complete, so we will not focus on that here.

Instead, we’re going to focus on how pterosaurs and bats got aloft and how this resulted in two radically different wing structures. By association, we will discuss how this relates to speculative evolution, as well as how to understand the mysterious volaticotheres.

High contrast


The basic bat and pterosaur wing models.

Although chiropteran and pterosaur wings are frequently compared, in truth they could be more functionally different.

The former evolved around the notion of grasping. Bat flight relies on the fingers bending as the wings move, an extension of the way mammal fingers normally flex (Cooper 2008, Swartz 1992).

To accomplish this task, bat wings are truly specialized hands, with all non-thumb fingers supporting the wing membrane. The digits themselves are elongated and highly flexible, individual bones even being able to bend considerably. Most interesting, most of the same tendon groups seen in other mammalian fingers remain, but they are thinned out to an extreme degree, hence the “skeletal” appearance of the wings.

Pterosaur wings, by contrast, are determined by the polar opposite, rigidity.


Pterosaur wing muscles.

Only one finger, the fourth digit, became a wing finger. Not only is this finger almost completely rigid – though some pterosaurs like anurognathids (Witton 2013) might have been able to bend the wing finger – its also oriented in a lateral way. It was unable to grasp the air.

Furthermore, the rest of the forelimb was also unable to grasp anything. All metacarpals are tightly pressed against each other, being unable to move significantly. The clawed fingers themselves have few phalanges (just one in the first digit, as in all sauropsids), rendering them rigid as well. The individual tendon systems associated with the digits also appear rather degenerate, except for the wing finger’s tendons, which are massive and would have made the hand area look rather fleshy in life.

Probably the best way to summarize this is to compare how both animals fold their wings:



Bats fold their wing fingers, flexing them like closed fists.

Hatz and Aram Witton 2015 low res

Azhdarchids by Mark Witton.

Pterosaurs, on the other hand, couldn’t do this, and their wing fingers stuck out like blades.

But why?


Iguana hand. Unlike mammals, which ancestrally have the same number of phalanges in all digits aside from the thumb, sauropsids have a gradient number of phalanges, from 1 in the first digit to as many as four in the fourth digit.

There are possibly various factors as to why we ended up with two wildly different wing types.

One could simply be the way sauropsid and synapsid hands are structured. While mammals have ancestrally the same number of phalanges in all digits (3) aside from the thumb (2), making them generally more evenly sized, in sauropsids the number of phalanges is gradient, increasing from the single phalange in the thumb, two in the second digit and so fourth until a maximum of five in the fifth (Witton 2013, Linzey 2012). Accordingly, in many reptiles like lizards this corresponds to a gradient in finger length.

To these ends, flexible, grasping hands are less common in sauropsids than they are in mammals. Most reptilian climbs prefer to use their digits as hooks, rather than grasp. Thus, a sauropsid flutterer might have been stressed to develop a more rigid wing, rather than a flexible one. The fact that rigid wings are present in pterosaurs, birds and scansoriopterygids would seem to prove my point.

However, I must admit that I’m not entirely satisfied with this explanation. A few sauropsids like chameleons have developed grasping hands (and, in the case of birds, grasping hindlimbs, which have the same phalangeal pressures as the forelimbs), so I think that, hypothetically, a reptile could develop grasping bat-like wings.

Rather, I think that the second possible factor is more likely. Simply, how bats and pterosaurs got airborne.

Bats, as mentioned before, became airborne by fluttering down. Baby bats still perform the hypothetical ancestral behavior: when approached from a high perch by a predator, they lunge themselves into the air, flapping their wings as to break the fall. Bat ancestors, likely arboreal or cave dwelling, likely flapped their forelimbs the same way. Boarder hands meant more efficient flapping, and over time this lead to wings.

Pterosaurs… are a total mystery. Its frankly impossible to say how exactly they became airborne.

However, the fact that non-pterodactyloid pterosaurs have hindlimbs specialized for hopping (Witton 2015), and that the hopping, kangaroo-like Scleromochlus is the closest known relative of pterosaurs (Witton 2013), it seems to suggest that pterosaurs evolved from a terrestrial hopping ancestor. This makes sense: at no point in pterosaur evolution did they ever develop arboreality, with even basal pterosaurs being more at home in the ground.

So, whatever process lead to pterosaur flight was likely ground focused, a true “ground up” scenario. My personal hypothesis is that pterosaur ancestors took the kangaroo-like hopping into a more lagomorph-like scenario, and the added boost provided by the forelimbs lead to flight. This hypothesis is based on the digitigrade nature of pterosaur forelimbs, which has been compared to ungulate mammals (Witton and Naish 2008).



Ichthyoconodon commission by Dylan Bajda. Here, the animal is depicted with styliform-based pterosaur-like wings.

Animal flight is often oversimplified. Its often interpreted as a sort of homogenous process, functionally identical in at least vertebrate animals, be them birds, pterosaurs or bats alike.

As such, its common in creature design to just depict it randomly when it comes to speculative evolution. Critters might have either bat or pterosaur-like wings at random, it being simply a matter of preference as far as the artist is concerned. I am myself guilty of this, having made up many flying animal clades in various projects.

However, it seems as though flight was unique in locomotion methods in that various groups developed it in radically different ways. While you can easily mistake the flippers and fins or whales, marine reptiles and fish or the digitigrade hindlimbs of various mammals and dinosaurs, the wings of pterosaurs, birds, bats and insects are nothing alike besides just being airfoils.

Structurally, they are very different organs impossible to be distinguished for one another. And as we find new winged animals like scansoriopterygids, we see that this uniqueness continues.

Which brings us to a group I’ve talked about a lot, the volaticothere eutriconodonts.

As I’ve mentioned before, there is evidence that these animals might have been capable of powered flight. And, to add to the ambiguity, the most complete specimen so far, the Volaticotherium antiquum, bears a rather incomplete hand (Meng 2006 additional materials), rendering the conversation all the more frustrating.

What little do we know suggests an odd forelimb. The metacarpals are similar to those of the hindfoot, but the proximal phalanges are longer than them and distally expanded. These details make the hands somewhat distinct from the feet, so while a conventional paw can’t be ruled out, it might also hint at a more unconventional forelimb.

Given the proposed grasping function of the forelimb phalanges, if it is a wing then it might have functioned somewhat like bat wings. However, unlike in bat wings the metacarpals were not significantly elongated, while the phalanges were. This would have implications both in functionality as well as how flight evolved.

If it was a more conventional paw, there’s the possibility that it had a long styliform element. Styliforms evolved multiple times in mammals and archosaurs (Xu 2015), so a wing structure dependent on styliforms, while speculative, is definitely plausible. If this was the case, then we might be looking at an entirely different way of acquiring powered flight.

Either situation would be a vindication of the established pattern: each flying vertebrate group is unique.


Glenn L. Jepsen, Bat origins and evolution, 1970

Cooper, K. L., & Tabin, C. J. (2008). Understanding of bat wing evolution takes flight. Genes & development, 22(2), 121-124.

Swartz, S. M., Bennett, M. B., & Carrier, D. R. (1992). Wing bone stresses in free flying bats and the evolution of skeletal design for flight. Nature, 359(6397), 726.

Witton, Mark P. (2015). “Were early pterosaurs inept terrestrial locomotors?”. PeerJ. 3: e1018. doi:10.7717/peerj.1018. PMC 4476129 Freely accessible. PMID 26157605.

Mark P. Witton (2013), Pterosaurs: Natural History, Evolution, Anatomy, Princeton University Press, ISBN 978-0-691-15061-1

Witton MP, Naish D (2008). McClain, Craig R., ed. “A reappraisal of azhdarchid pterosaur functional morphology and paleoecology”. PLoS ONE. 3 (5): e2271. doi:10.1371/journal.pone.0002271. PMC 2386974 Freely accessible. PMID 18509539.

Donald W. Linzey, Vertebrate Biology, JHU Press, 2012

Meng, J.; Hu, Y.; Wang, Y.; Wang, X.; Li, C. (2006). “A Mesozoic gliding mammal from northeastern China”. Nature. 444: 889–893. doi:10.1038/nature05234.; specifically, the associated appendix

Xu, X.; Zheng, X.; Sullivan, C.; Wang, X.; Xing, L.; Wang, Y.; Zhang, X.; o’Connor, J. K.; Zhang, F.; Pan, Y. (2015). “A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings”. Nature. 521: 70–3. doi:10.1038/nature14423. PMID 25924069.

Basic rundown of Choristodera, the “freshwater marine reptiles”

April 24, 2017

a) Simoedosaurus lemoinei (modified from Sigogneau-Russell and Russell, 1978) –b) Ikechosaurus pijiagouensis (IVPP V13283) –c) Tchoiria namsarai (redrawn from Efimov, 1975) –d) Champsosaurus gigas (SMM P77.33.24): non-neochoristoderan –e) Cteniogenys sp. (Evans, 1990)–f) Lazarussuchus inexpectatus (Hecht, 1992) –g) Hyphalosaurus baitaigouensis (redrawn from Gao and Ksepka, 2008) –h) Monjurosuchus splendens (Matsumoto et al., 2007) –i) Philydrosaurus proseilus (Gao and Fox, 2005).

Choristoderes are a group of extinct reptiles I’ve talked about before. Now that several years have passed, I’m going to discuss them once more.

What are choristoderes?


Hyphalosaurid choristoderes (and Manchurochelys) by kahless28

Choristoderes are a lineage of extinct, generally fully-aquatic reptiles. Beyond that, however, it’s very hard to tell.

See, it’s been a long standing debate whereas choristoderes are either archosauromorphs (the lineage of reptiles leading to archosaurs, including crocodylomorphs, pterosaurs and dinosaurs), lepidosauromorphs (the lineage leading to squamates and sphenodontians), or neither (more basal than either group). Similar debates have also plagued classical marine reptile groups like plesiosaurs and ichthyosaurs.

Part of the reason why figuring out where choristoderes fit in the phylogenetic tree of reptiles is because they lack definitive features affiliating them to either major reptile group. Their skulls vaguely resemble those of lepidosauromorphs, but lack the same quadrate configuration.

A particularly radical proposition was suggested by Miller 2004, in which choristoderes are linked to classical marine reptiles (ichthyosaurs and sauropterygians) under Euryapsida.

Personally, I lean slightly towards a lepidosaur identity, but further results may prove this hunch wrong.


When did they live?


Hyphalosaurus by Matt Martuniuk.

Pachystropheus and Actiosaurus might represent examples of Rhaetian Triassic choristoderes. There is still some ambiguity; the former has also been regarded as a sauropterygian (Matsumoto 2009) or as a thalattosaur (Renesto 2005), while the latter was initially interpreted as a theropod, then an ichthyosaur before its proposed choristodere identity more recently (Mortimer 2010).

If both or at least one are choristoderes, they’re both fairly specialized marine reptiles and, in Pachystropheus‘ case, fairly derived, implying an even earlier origin. This is consistent with the long temporal gaps in choristodere fossil record history, the Lazarus taxon effect.

Otherwise, choristoderes are first known from the mid/late Jurassic. These Jurassic species are less specialized, being lizard-like freshwater reptiles. This would set a norm for the rest of the group’s existence: highly conservative “living fossils” whose lineages bear a spotty fossil record, attesting to both their resilience as well as avoidance of preservation. These include the complicated “Cteniogenys” assemblage, which first arose in the mid-Jurassic of Europe and North America, including the famous Morrison Formation.

By the Early Cretaceous, choristodere diversity in Asia undergoes an explosive radiation. Besides the more classical, salamander-like monjurosuchids, we also see the arrival of the long-necked hyphalosaurids and the long-jawed neochoristoderes, expanding the ecological niches available to the group.

Its unclear why choristoderes underwent such a sudden diversification. The local extinction of aquatic crocodilians due to colder temperatures might have triggered the evolution of neochoristoderes (Matsumoto 2010), but this doesn’t explain the evolution of hyphalosaurids.

From the mid-Cretaceous onward, there are no Asian choristodere remains, other than the largely understudied Eotomistoma (Carroll 1988). Hyphalosaurids disappear, but North America and Europe see the presence of both salamander-like species (“Cteniogenys”) as well as the diverse neochoristodere Champsosaurus. Both groups co-exist with a variety of aquatic crocodilians, though Champsosaurus apparently prevents long-snouted crocodilians from occurring in freshwater habitats (Matsumoto 2010).

Choristoderes survived the KT event, with Champsosaurus displaying dietary changes (Matsumoto 2015). It undergoes an adaptive radiation, producing several species including C. gigas, the largest known choristodere at around 3.5 meters long. It is joined by another genus, Simoedosaurus, which given its sudden appearance in the fossil record probably evolved from a Cretaceous Asian species.

Cretaceous “Cteniogenys” species are not known to have made it past the KT event. However, the genus Lazarussuchus debuts in the Paleocene of France, a much more basal animal with a ghost lineage quite possibly extending to the Jurassic or Triassic (Hecht 1992), unless they are related to the Cretaceous “Cteniogenys” species (Matsumoto 2013).

Neochoristoderes suddenly disappear in the medial Eocene, for no currently explainable reasons (see below). However, Lazarussuchus would continue to endure until the mid-Miocene, if not possibly until the Pliocene, when Europe grew too cold for these reptiles.

Unlike many clades, in which the most specialized members were the last representatives, choristoderes seem to have ended more or less as they started.

Fully Aquatic


Hyphalosaurus giving birth by Joschua Knüppe.

Most choristoderes were fully aquatic animals.

While some basal taxa like Lazarussuchus could be interpreted as amphibious, both monjurosuchids and hyphalosaurids have evidence of vivipary (Wu 2010) while in Champsosaurus only adult females have limbs robust enough to carry them ashore, while males and juveniles could not support their weight on land (Katsura 2007). As such, choristoderes as a whole appear to not have left the water much, if at all.

All known choristoderes possess laterally flatted tails, which were probably their main propulsion mechanism, with neochoristoderes also having paddle-like limbs. At least hyphalosaurids, monjurosuchids and neochoristoderes all have smooth skins with small, non-overlapping scales, though monjurosuchids do bear rows of scutes similar to those of modern alligator-lizards (Gao 2000). In neochoristoderes, the torso is dorsoventrally flattened, the gastralia are large and the ribs are short and massive, further adaptations for diving. There was a source mentioning pachyostic bones, but I can’t seem to find it.

Probably the most remarkable adaptation for life in the water are the nostrils. Aside from basal choristoderes like Lazarussuchus, which simply have receded nostrils, in more derived taxa the nostrils are fused and oriented towards the tip of the snout, allowing the animals to effectively use them as a snorkel, surfacing only the very tip of the snout while the rest of the body remained underwater (Acorn 2007).

Perhaps tellingly, the eyes are forwardly oriented. This is in contrast to amphibious animals like crocodilians and hippos, which have dorsally-oriented nostrils and eyes.

Cold Waters


Palaeogeographical distribution of Choristodera in the Jurassic and Cretaceous periods.

Choristoderes as a whole have a pan-Laurasian distribution, ocurring in fossil sites in North America, Europe and Asia. There are two possible exceptions: basal choristoderes from the Jurassic/Early Cretaceous of North Africa (Haddoumi 2014) and possible neochoristodere teeth from Timor (Umbgrove 1949).

Other than these examples, however, nearly all choristodere remains occur in high latitudes. Although some do occur in sites that had a paratropical climate, most occur in temperate regions. Some Champsosaurus fossils even ocur in the high Arctic, in the Cretaceous of Greenland and the Eocene and Cretaceous of Axel Heiberg (Matsumoto 2010).

Thus, it can be seen that choristoderes tolerated colder environments, if not outright preferred them (Matsumoto 2014). Their ability to survive in colder climates in fact probably allowed them to diversify in the absence of crocodilians during the Early Cretaceous, when colder temperatures in Asia caused the local extinction of aquatic crocodilians (Matsumoto 2014). Both groups ultimately co-existed in warmer areas, however.

I previously speculated that neochoristoderes might have been endothermic, which fits well with their extreme speciation to an aquatic lifestyle. However, given that cold-tolerance was also present in more basal taxa, monjurosuchids and hyphalosaurids, I wonder if it endothermy was more widespread across the clade.

Ironically, cooling temperatures in Europe probably lead to the extinction of the group, as Lazarusussuchus probably survived until the glaciations.




Palatine teeth in Simoedosaurus.

Basal choristoderes were probably generalistic feeders, ambushing small fish, crustaceans and other prey. Monjurosuchus has evidence of arthropod cuticle in its intestines, suggesting that it fed on aquatic arthropods (Gao 2000).

Hyphalosaurids were certainly specialized animals, bearing long necks. It’s likely that they were doing whatever the convergently similar plesiosaurs were doing, whatever that was. In particular, they appear to have preferred softer prey, which they probably hunted actively on the deeper areas of their lake environments (Gao 2008).

By contrast, the long-snouted neochoristoderes are much easier to figure out. These predators were doing what crocodilians, whales, gars and countless other large, predatory aquatic vertebrates do best: snatch up larger prey. Long, thinner jaws allowed them to accomplish this task with less drag.

While they are often compared to gharials, neochoristoderes may not be exactly analogous. Unlike other crocodilians, gharials have weak bite forces, rarely exceeding 497 N; they can tackle animals as large as goats, but they prefer small fish. By contrast, Champsosaurus had a bite force of 1194 to 1910 N, more comparable to that of crocodiles (James 2010). This may not have necessarily translated to a preference for larger prey, but it does convey a much larger degree of strength and speed when catching small fish.

Simoedosaurus dakotensis and Champsosaurus gigas, by contrast, have semi-brevirostrine snouts, and might have a diet more similar to that of crocodiles, feeding on not just fish but aquatic tetrapods like turtles and waterfowl and maybe ambushing terrestrial animals near the water. Notably, they co-existed with crocodile-like crocodilians like Borealosuchus (Matsumoto 2013 and 2014).

A neat feature choristoderes have is the retention of palatine teeth. Lost in various tetrapod groups like mammals, crocodylomorphs and dinosaurs, these teeth basically serve the role palatal grooves have in these groups, helping the animal to hold and manipulate food items in the mouth. In basal choristoderes these teeth are fairly generic, while in neochoristoderes they are more specialized, indicating a higher degree of food manipulation in the mouth. Such speciation might have evolved multiple times, as the simoedosaurid Ikechosaurus has palatine teeth more similar to those of non-neochoristodere choristoderes (Matsumoto 2015).

Through subtle differences in palatine teeth, we know that Champsosaurus changed its diet across the KT event, and that Simoedosaurus lindoei prefered softer prey than S. dakotensis (Matsumoto 2015).



Lazarrusuchus, the last choristodere, by Nobu Tamura.

As mentioned previously, the extinction of choristoderes isn’t clear.

Competition with crocodilians is sometimes attributed to the demise of neochoristoderes, but both groups co-existed across the Late Cretaceous, Paleocene and Eocene of Laurasia and possibly Timor, and if anything neochoristoderes seem to have been dominant over long-snouted crocodilians in freshwater habitats (Matsumoto 2010 & 2014).

This also doesn’t explain the demise of hyphalosaurids, or of basal choristoderes in North America, North Africa and Asia.

Ultimately, European choristoderes such as Lazarussuchus appear to have died out due to glaciations in Europe.


There is still much I haven’t covered. Hopefully this will introduce you to these amazing extinct reptiles.


Ryoko Matsumoto; Shigeru Suzuki; Khisigjav Tsogtbaatar; Susan E. Evans (2009). “New material of the enigmatic reptile Khurendukhosaurus (Diapsida: Choristodera) from Mongolia”. Naturwissenschaften. 96 (2): 233–242. doi:10.1007/s00114-008-0469-6. PMID 19034405.

Silvio Renesto (2005). “A possible find of Endennasaurus (Reptilia, Thalattosauria) with a comparison between Endennasaurus and Pachystropheus“. Neues Jahrbuch für Geologie und Paläontologie – Monatshefte. Jg. 2005 (2): 118–128.

R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274

R. L. Carroll. 1988. Vertebrate Paleontology and Evolution. W. H. Freeman and Company, New York 1-698

R. Matsumoto and S. E. Evans. 2015. Morphology and function of the palatal dentition in Choristodera Article in Journal of Anatomy 228(3):n/a-n/a · November 2015 DOI: 10.1111/joa.12414

Hecht, M.K. (1992). “A new choristodere (Reptilia, Diapsida) from the Oligocene of France: an example of the Lazarus effect”. Geobios. 25: 115–131. doi:10.1016/S0016-6995(09)90041-9.

Matsumoto, R.; Buffetaut, E.; Escuillie, F.; Hervet, S.; Evans, S. E. (2013). “New material of the choristodere Lazarussuchus(Diapsida, Choristodera) from the Paleocene of France”. Journal of Vertebrate Paleontology. 33 (2): 319. doi:10.1080/02724634.2012.716274.

Ji Q., Wu, X.-C. and Cheng, Y.-N. (2010). “Cretaceous choristoderan reptiles gave birth to live young.” Naturwissenschaften, 97(4): 423-428. doi:10.1007/s00114-010-0654-2

Yoshihiro Katsura. 2007. Fusion of sacrals and anatomy in Champsosaurus (Diapsida, Choristodera), doi:10.1080/08912960701374659

Gao, K.; Evans, S.; Ji, Q.; Norell, M.; Ji, S. (2000). “Exceptional fossil material of a semi-aquatic reptile from China: the resolution of an enigma”. Journal of Vertebrate Paleontology. 20 (3): 417–421. doi:10.1671/0272-4634(2000)020[0417:efmoas];2.

John Acorn, Deep Alberta: Fossil Facts and Dinosaur Digs, University of Alberta, 07/02/2007

Hamid Haddoumi, Ronan Allain, Said Meslouh, Grégoire Metais, Michel Monbaron, Denise Pons, Jean-Claude Rage, Romain Vullo, Samir Zouhri, Emmanuel Gheerbrant, Guelb el Ahmar (Bathonian, Anoual Syncline, eastern Morocco): First continental flora and fauna including mammals from the Middle Jurassic of Africa, doi:10.1016/

J. H. F. Umbgrove, Structural History Of The East Indies

Gao, K.-Q. and Ksepka, D.T. (2008). “Osteology and taxonomic revision of Hyphalosaurus (Diapsida: Choristodera) from the Lower Cretaceous of Liaoning, China.” Journal of Anatomy, 212(6): 747–768. doi:10.1111/j.1469-7580.2008.00907.x

James, Michael, The jaw adductor muscles in Champsosaurus and their implications for feeding mechanics, 2010-08-30T19:01:39Z

Six Reasons Why Eutriconodonts Are Awesome

April 22, 2017

People who know will definitely tell you how much I’ve come to appreciate Mesozoic mammals. Usually dismissed as small rat things, mammals in the Mesozoic were a highly diverse bunch of animals, including swimmers, diggers, anteater like forms, large terrestrial predators, hoppers and many, many more.

Of these, eutriconodonts are by far among the more spectacular. I’ve already talked at length about the possible flight capacities of volaticotheres, but really the whole clade is pretty neat, and here’s why:

1- The first mammalian carnivores

Jugulator amplissimus by @paleoart

Eutriconodonts are notable for being among the first mammals specialized to dedicated carnivory. Zofia Kielan-Jaworowska identified numerous features associated with obligate carnivory: long, sharp canines (or canine-like incisors in the case of gobiconodontids), premolars with trenchant main cusps that were well suited to grasp and pierce prey, strong development of the mandibular abductor musculature, bone crushing ability in at least some species and several other features.

Their iconic triconodont dentition, usually taken as “primitive”, might actually be specialized for shearing (Zofia Kielan-Jaworowska 2004, Sigogneau-Russell 2016), making it vaguely analogous to the carnassials of placentals and marsupial predators. Their exact shearing mechanism has no real analogue among mammalian carnivores, but the function is considered very similar at least (Rougier 2015)

Equally important is eutriconodont size. Eutriconodonts are among the largest mammals in Mesozoic faunal communities, which has been inferred as standing the highest among mammals in contemporary trophic webs ( Zofia Kielan-Jaworowska 2004). At their size, they were perfectly capable of taking down vertebrate prey, and the largest gobiconodontids like the infamous Repenomamus might have been apex predators in their environment.

Repenomamus itself has been found with dinosaur remains in its belly, and scavenging marks associated with Gobiconodon have also been found. These mammals could, in fact, tackle dinosaurs, and if modern analogues like wolverines, tasmanian devils and ratels are of any indication then the largest eutriconodonts could in fact be “top guns” in their environments.

Other Mesozoic synapsids have also been inferred to be specialized carnivores, like Sinoconodon and deltatheroideans. But they lived either before eutriconodonts spread, or after they became extinct, and as such their range was much more limited.

2- Their diversity

Speculative depictions of

Ichthyoconodon by @alphynix. While I argue for a slightly different lifestyle, they help summarize the range of known eutriconodont bauplans, such as the otter like Yanoconodon and Liaoconodon and the aerial Volaticotherium and Argentoconodon

Better only than carnivorous Mesozoic mammals are carnivorous Mesozoic mammals that come in all shapes and sizes. In spite of being pretty much highly specialized carnivores and certainly more restricted in terms of diet than, say, symmetrodonts or early therians, eutriconodonts were much more diverse than these groups were (until therians got them beat after eutriconodonts went extinct, that is).

The group ranged from shrew analogues (amphilestids, amphiodontids, basal gobiconodonts and some triconodontids), arboreal, tree-shrew like forms (Jeholodens), large, robust carnivores (gobiconodontids, Jugulator, triconodontids), a quilled species with an immensely thick spine (Spinolestes), at least two lineages of swimmers (Liaoconodon and Yanoconodon) and of course the aforementioned volaticotheres, conservatively gliders if not outright flyers.

The exact smallest triconodonts probably weighted around 50 grams. The largest, Repenomamus giganticus, as much as 14 kg.

This level of ecological diversity is so far unmatched by any Mesozoic mammal group save for multituberculates and perhaps Late Cretaceous metatherians. It is even larger than the diversity of most therian carnivore groups, save for carnivorans.

Every possible niche taken by carnivorous mammals under 14 kg was taken, and it’s amazing.

3- Brains!

Yes, we know about eutriconodont brains. In fact, Triconodon mordax is one of the first extinct animals to have its endocast studied (Simpson 1928).

From what we can tell, at least from this one specimen, eutriconodonts had fairly “primitive” brains for mammal standards. The cerebral hemisphere is long, oval and flat, lacking the inflated appearance present in modern mammals (including monotremes, which are generally held to be more basal than eutriconodonts!) as well as the also extinct multituberculates. The cerebrum is similarly not expanded as much as in those groups. Like multies, Triconodon has a large, semi-triangular bulge, thought to be a large cistern.

What this means about eutriconodont intelligence is unclear. It might seem like they were fairly stupid mammals, but mammals with fairly simplistic brains are known to be fairly intelligent (Weisbecker 2010). They probably weren’t as cunning as modern cats and dogs, but probably capable of complex behaviors nonetheless.

4- Everywhere For A Long Time

Different mammaliaform tooth types across the Mesozoic. Eutriconodonts, alongside the unrelated morganuconodonts, were the only mammals to bear a “triconodont” tooth type.

While eutriconodonts fall short of multituberculates as the longest living mammal lineage, they were still very successful. The first eutriconodont fossils – Argentoconodon, Victoriaconodon and Huasteconodon – all date to the Toarcian and represent a large variety of lineages, indicating an even earlier origin.

Eutriconodonts would then keep on going in full force for another 111 million years. Even when other mammal groups display gaps in their fossil record, eutriconodonts continue across fossil sites in Europe, Asia, North America, Africa and South America, rendering them a truly global presence in Mesozoic faunas as much as dinosaurs and pterosaurs.

Alas, they faced a final challenge with the spread of angiosperm plants, which drastically altered faunal components across the globe and was particular harsh on carnivorous mammals. Only one lineage survived the Turonian, Alticonodon, to still endured all the way to the Campanian.

5- Poison-Heels


by Dmitry Bogdanov. Notice the spurs on the heels.

Okay, not something exclusive to eutriconodonts among mammals, but it bears repeating.

Venomosity is inferred to be an ancestral trait for mammals (Hurum 2006). Various Mesozoic mammal groups possess heel spurs similar to those of the modern platypus, which delivers a powerful neurotoxin infamous for how painful it is. This includes similar canals, which implies an identical function.

This spurs have been found in nearly all non-therian mammal groups, suggesting that either venom evolved multiple times among Mesozoic mammals, or, most likely, that it was an ancestral feature later lost in therians.

Eutriconodonts, of course, preserve such spurs. They are best known in gobiconodontids, which combined with other features would make these some of the most ridiculously over-engineered killing machines of the time.

6- Tough As Nails

Fantastic Mr. Spinolestes.

Gobiconodontids were probably the honeybadgers of the Jurassic (and early Cretaceous). The largest of all eutriconodonts, they included not only the infamous Repenomamus, but the also fairly sizeable Gobiconodon. These animals are racoon-to-wolverine sized beasts, bearing thick skeletons, robust jaws and sharp fang-like incisors.

As such, not only were they large carnivorous mammals for the time, but also specifically designed to fight violently. Combined for evidence for scavenging for Gobiconodon and outright dinosaur-consumption for Repenomamus, it’s hard to not see these as competitors for small to mid-sized theropod dinosaurs in their local environments. “Small”, but incredibly brute fighters, fighting their way into carcasses and perhaps even harassing fellow predators.

That said, even the smaller gobiconodontids were nothing to laugh at.

Spinolestes, a more conventionally shrew-sized animal, bears:

– A massively thick, xenarthrous spine similar to that of xenarthrans and the hero shrew. This probably allowed it to survive being smashed by animals up to 75 kg.

– Spines similar to those of the modern spiny mice.

– The venom spurs.

What can you even say to that?


I think Mesozoic mammals are underrated in general, but eutriconodonts in particular are a very fascinating group. Besides these undeniably awesome facts, there’s also the fact that they bear some of the most exquisitely preserved Mesozoic mammal fossils, something even the more well known multituberculates currently lack.

Dismissed as just archaic “missing links”, they were a dynamic, fascinating group of animals, which I believe deserve some recognition.


Zofia Kielan-Jaworowska, Richard L. Cifelli, Zhe-Xi Luo (2004). “Chapter 7: Eutriconodontans”. Mammals from the Age of Dinosaurs: origins, evolution, and structure. New York: Columbia University Press. pp. 216–248. ISBN 0-231-11918-6.

Percy M. Butler; Denise Sigogneau-Russell (2016). “Diversity of triconodonts in the Middle Jurassic of Great Britain” (PDF). Palaeontologia Polonica 67: 35–65. doi:10.4202/pp.2016.67_035.

Chen, Meng; Wilson, Gregory P. (2015). “A multivariate approach to infer locomotor modes in Mesozoic mammals”. Paleobiology. 41 (02): 280–312. doi:10.1017/pab.2014.14. ISSN 0094-8373.

Vera Weisbecker and  Anjali Goswami, Brain size, life history, and metabolism at the marsupial/placental dichotomy, Proc Natl Acad Sci U S A. 2010 Sep 14; 107(37): 16216–16221. Published online 2010 Sep 7.   doi:  10.1073/pnas.0906486107

Flying Volaticothere Developments

February 27, 2017




volbatUpdated depictions of Argentoconodon, Triconolestes and Ichthyoconodon (Ceri Thomas and Dylan Bajda, respectively), and a fictional species for the Speculative Dinosaur Project by Tim Morris.

So a while ago I made some  posts on whereas a group of Mesozoic mammals, the volaticotheres, were capable of true powered flight. There have been relatively few responses since then, other than the expected “take this with a grain of salt” (which is fair; it’s an outrageous hypothesis), and even fewer addressals to the points I provided. I had a conversation with paleofail-explained on Julio Lacerda’s commission, but it was abandoned quickly. A more successful counter-argument was made by fezraptor, which I will address below.

In the past few months there have been relatively few papers involving eutriconodont mammals, and none of them involving volaticothere taxa. Thus, for now, things remain as they remain.

However, I previously did miss an important study, which deals with the dietary ranges of Mesozoic mammals and their relation to angiosperm diversity. Here, both Argentoconodon and Volaticotherium are present, firmly within the animalivorous range, with the former ranking more with carnivorous taxa and the latter with insectivorous taxa. Animalivorous mammals are shown to have declined substantially during the mid-Cretaceous; both points strengthen the arguments for volaticotheres being specialised animalivores, and thus unlikely to have been gliders.

Now, unto for the addressal:

The manus of Volaticotherium is actually sufficiently preserved to show that it is smaller than the pes (barring the presence of unprecedentedly weird distal phalanges), precluding it looking bat-like.

The appendix for Meng 2003 describes the manus as “poorly preserved”, and doesn’t mention size. Granted, the fact that the metacarpals don’t seem to be particularly specialised in relation to those in the foot makes a bat-like wing less likely, but the aforementioned elongated phalanges might suggest an atypical wing structure.

Ichthyoconodon is actually outside of the clade that preserves any evidence of adaptations for gliding, so kenbrasai’s reasons for why Ichthyoconodon would be atypical for a glider actually strike me as evidence it wasn’t volant at all.

True, though it’s worth to note that Ichthyoconodon‘s exclusive status from this clade is based on one character less (Gaetano et al 2011). In any case Ichthyconodon‘s molars are less recurved than those in Volaticotherium and Argentoconodon, and as these taxa are already disparate in their jaw morphology it could mean that Ichthyoconodon was functionally very different. We do have evidence of truly aquatic eutriconodonts, after all.

I do stand by my hypothesis, however, as it is still a rare taxon, so an aquatic lifestyle seems less likely.