Roman Uchytel. Taking paleoart to the next level.

Figure 1. Roman Uchytel is the arist/naturalist who is bringing prehistoric beasts and birds back to life.

Figure 1. Roman Uchytel is the arist/naturalist who is bringing prehistoric beasts and birds back to life.

Here’s an artist worth noting.
Roman Uchytel (Fig. 1) says it best himself, “Using only their skeletons, I bring creatures to life that roamed the same routes that take you to and from work hundreds of thousands of years ago.”

His mission:
“Roman Uchytel’s galleries constitute the first resource solely dedicated to the reconstruction of prehistoric animals beyond the dinosaurs. These are not photographs, but rather, artistic recreations from the skeletons of ancient animals that roamed the earth millions of years ago. Many of these fascinating creatures are unfamiliar to the public and remain a mystery even to science.”

Figure 2. Homepage for Roman Uchytel's images. Click to visit.

Figure 2. Homepage for Roman Uchytel’s images. Click to visit.

Check out his website
and you will be filled with wonder: https://prehistoric-fauna.com

 

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What is Fruitafossor? A xenarthran close to Peltephilus.

This appears to be
yet another case of pertinent taxon exclusion. Today’s fossorial digger has several universally acknowledged xenarthran (edentate) traits. For reasons unknown it was not tested against another fossorial xenarthran, Peltephilus. Rather the authors compared their digger to an arboreal sloth, Bradypus, among several other taxa, including distinctly different anteaters and armadillos.

Figure 1. Scapula of Fruitafossor compared to several candidate sisters. Luo and Wible made things a bit more difficult by presenting left and right scapulae. Here they are all left scapulae for ready comparison. There is no doubt that the Fruitafossor scapula looks more like that of Ornithorhynchus.

ºªº Figure 1. Scapula of Fruitafossor compared to several candidate sisters. Luo and Wible made things a bit more difficult by presenting left and right scapulae. In frame 2 they are all left scapulae for ready comparison. There is no doubt that the Fruitafossor scapula was illustrated to look more like that of Ornithorhynchus. Unfortunately the photo data (Fig. 2) does not clearly support that shape. That shape is so important, it needed to be better documented.

Luo and Wible 2005
brought us a small, mostly articulated, rather crushed and incomplete Late Jurassic mammal with simple blunt teeth and digging forelimbs. Fruitafossor windscheffeli (Figs. 1–6) is best represented by a CT scan (Figs. 2–4) and original drawings (Figs. 5, 6) created by the Luo and Wible team.

Figure 2. Fruitafossor in situ from Digimorph.org and used with permission and here colorized to an uncertain extent.

Figure 2. Fruitafossor in situ from Digimorph.org and used with permission and here colorized to an uncertain extent. All those little white dots could be scattered osteoderms.

The original analysis
nested Fruitafossor between extremely tiny Hadrocodium + Shuotherium and Gobiconodon in a tree topology that does not resemble the topology of the large reptile tree (LRT, 1048 taxa). The authors noted Fruitafossor is “not a eutherian, let alone a xenarthran” despite noting Fruitafossor had tubular molars and xenarthran intervertebral articulations, traits otherwise found only in xenarthrans.

Figure 2. Same specimen from Digimorph.org and rotated to show the teeth better. See figure 3 for a closeup.

Figure 3. Same specimen from Digimorph.org and rotated to show the teeth better. See figure 3 for a closeup. All those little white dots could be scattered osteoderms. Some of this flatness is due to crushing. Some of it is due to this being a wider than deep armored mammal.

Wider than deep
Yes, the Fruitafossor specimen is crushed, but what is shown here indicates a low, wide mammal just getting some armor in the Late Jurassic. And we all know why armor might have been helpful! And this may explain the lateral sprawl of the forelimbs and giant wide humerus, another atavism!

Figure 3. Closeup of figure 2 showing maxilla and dentary in occlusion.

Figure 4. Closeup of figure 2 showing maxilla and dentary in occlusion. If there is a convex ventral dentary it must be imagined because it is not preserved.

By contrast,
the LRT nested Fruitafossor with the horned, armored digging ‘armadillo’ more closely related to BradypusPeltephilus (Fig. 5).

Figure 4. The xenarthran, Peltephilus, compared to Fruitafossor, not to scale, but to similar jaw lengths. Note the drawing from Zhou and Wible does not exactly match what one can see in the photo (Fig. 3). This data needs to be clear and it is not.

Figure 5. The xenarthran, Peltephilus, compared to Fruitafossor, not to scale, but to similar jaw lengths. Note the drawing from Luo and Wible does not exactly match what one can see in the photo (Fig. 3). This data needs to be clear and it is not.

Luo and Wible compared
Fruitafossor to the arboreal and extant Bradypus, but not to the fossorial and extinct Peltephilius (Fig. 5). I would consider that a mistake or an oversight that here overturns their hypothesis of a relationship of Fruitafossor to basalmost mammals.

Figure 5. Several drawings from Zhou and Wible that one must trust for accuracy. The verification data is too fuzzy to validate.

Figure 6. Several drawings from Luo and Wible that one must trust for accuracy. The verification data is too fuzzy to validate. As in other xenarthrans, the ilia actually form a pair of horizontal plates on either side of the long fused and eroded sacrals. Four fingers is a trait shared with Peltephilus. Imagine that rib cage wider and not so deep.

Scattered osteoderms
(Fig. 3) were not mentioned in the text. That’s one more trait shared with the armored xenarthran, Peltephilus. This overlooked relationship of derived xenarthrans moves them into the Jurassic, an era they have never been in before in phylogenetic and chronological analyses. Here placental arboreal and fossorial mammals (prior to condylarths) shared time and space with dinosaurs during the Jurassic and Cretaceous, but have, so far, been underrepresented in the fossil record. That’s changing with re-examination of the data applied to a larger gamut taxon list.

Although the illustrated scapula
(Fig. 1) looks like that of an egg-laying mammal, I will wait for better data to validate that illustration. In the meantime, Fruitafossor has blunt, tubular molars and xenarthran vertebral articulations (among many other xenarthran traits) because it is a xenarthran, not an egg-layer with convergent traits.

References
Luo Z-X and Wible JR 2005. A late Jurassic digging mammal and early mammal diversification. Science 308:103–107.

Vilevolodon: the atavistic reappearance of post-dentary bones

Preface
I’ve been wondering about the traditional nesting of Multituberculata and kin outside of the Mammalia for years. All have a dentary jaw joint, but some have post-dentary bones. given the opportunity multituberculates nest with rodents and plesiadapiformes in the large reptile tree (LRT, 1047 taxa).  No other pre-mammals resemble them. Traditionally Haramiyava (Fig. 1) has been considered a pre-mammal link to Haramiyida + Multituberculata. In the LRT Haramiyava nests with the mammaliaforms Brasilodon, Sinoconodon and Therioherpeton – far from any other taxa considered Haramiyida + Multituberculata currently and provisionally nesting deep within the Mammalia.

Figure 1. Haramiyavia reconstructed and restored. Missing parts are ghosted. The fourth maxillary tooth appears to be a small canine. The post-dentary bones are imagined from Vilevolodon (figure 4).

Vilevolodon diplomylos
(Luo et al. 2017; Jurassic, 160 mya; BMNH2942A, B; Figs. 2-4) was originally considered a stem mammal (= mammaliaform), a eleutherodontid in the clade Haramiyida AND it had clearly defined gliding membranes (Fig. 2). By contrast the LRT nests Vilevolodon with the Late Jurassic para-rodent Shenshou and the extant rodents, Rattus and Mus, not far from members of the Multituberculata.

Figure 1. Vilevolodon in situ, plate, counterplate, original drawing, DGS color, and restored manus and pes. Note the gliding membrane (patagium) and fur.

Figure 2. Vilevolodon in situ, plate, counterplate, original drawing, DGS color, and restored manus and pes. Note the gliding membrane (patagium) and fur.

But there’s a big problem
Vilevolodon doesn’t have tiny ear bones, like mammals do. It has post-dentary bones, like pre-mammals do (Figs. 3, 4).

Figure 2. Vilevolodont skull in situ, without color, DGS color tracing, that tracing reconstructed and a CT scan form Luo et al. 2017.

Figure 3. Vilevolodont skull in situ, without color, DGS color tracing, that tracing reconstructed and a CT scan form Luo et al. 2017.

The ear problem in Jurassic rodents
Luo et al. report, “a mandibular middle ear with a unique character combination previously unknown in mammaliaforms.” Pre-mammals have post-dentary bones (articular, angular, surangular). Therian mammals shrink and migrate those bones to the base of the skull where they become middle ear bones with new names (malleus, incus, ectotympanic). The stapes remains the stapes in all tetrapods. So what is happening with Vilevolodon and its sisters? Why don’t the pre-mammal post-dentary bones define it as a pre-mammal? After all, that’s the current paradigm.

Figure 3. There is no doubt that Vilevolodon has a pre-mammal type of posterior jaw bones. Otherwise they nest with rodents and plesiadapiformes. This appears to be a mammal with an atavism, a reversal. These elements simply stopped developing as in other mammals.

Figure 3. There is no doubt that Vilevolodon has pre-mammal type post-dentary bones. There is also no doubt that the dentary formed the main jaw joint with the squamosal. How does one reconcile both sets of traits? In the LRT Vilevolodon nests with rodents. This appears to be a mammal with an atavism, a reversal. These elements simply stopped developing as in other mammals.

Mammals are defined by
the evolution and migration of their posterior jaw bones into middle ear bones with a jaw joint switch from quadrate/articular to dentary/squamosal. Multituberculates and haramiyids appear to bend or break that rule because they have cynodont-like posterior jaw bones, not tiny middle ear bones, and yet otherwise they nest with rodents and plesiadapiformes. This is one reason why you don’t want to pull a Larry Martin with post-dentary bones. You want to nest a taxon based on a long list of traits, not just one, two or a dozen.

The massive jaw joint
Mammals, such as Vilevolodon, with atavistic post-dentary bones also have a massive jaw joint with a long articulating surface on the dentary contacting the squamosal. All mammals have such a jaw joint. Pre-mammals don’t. While Vilevolodon has a large dentary/squamosal jaw joint, the post-dentary articular, still contacts the quadrate. It’s clearly not the main jaw joint.

Filan 1991
traced the development of post-dentary bones in embryonic Monodelphis specimens. She reported, “Neonates of Monodelphis possess neither mammalian (dentarysquamosal) nor reptilian (quadrate-articular) jaw articulations, nor does the contact between the incus and crista parotica offer a joint surface. Elasticity in Meckel’s cartilage allows minimal deflection of the lower jaw.” After all, those neonates are just sucking milk, not biting, and the embryos don’t even do that. Does that make neonates like this not mammals? No. The evidence indicates that in multituberculates and haramiyds the embryological transformation of posterior jaw bones stopped before development transformed them into middle ear bones. This is an atavism, a phylogenetic reversal. The timing of development changed. In the case of Vilevolodon, the middle ear bones stop evolving during embryological development and the post-dentary bones they would have evolved from continue to appear in adults. What was a rare mutation probably spread throughout an isolated population. Perhaps this had something to do with the increase in size of the dentary jaw joint.

Haramiyavia and the Haramiyida clade
Seems at this point that only Haramiyavia is a haramiyid, unless Brasilodon is one as well. Members traditionally assigned to the clade Eleutherodontidae also nest in various locations in the LRT, not all in one clade.

Meng et al. 2017 report,
“Stem mammaliaforms are morphologically disparate and ecologically diverse in their own right, and they developed versatile locomotor modes that include arboreal, semiaquatic, and subterranean specializations, which are all distinct from generalized mammaliaforms.” Unfortunately, the LRT nests a long list of mammaliaforms at various nodes within the Mammalia. They are not from a single diverse clade.

Contra Meng et al. 2017
the LRT reduces the niches and body shapes of stem mammals down to a few small, generalized taxa like Sinoconodon and Megazostrodon. Derived taxa nest at derived nodes.

The LRT nests rodents close to Plesidapiformes,
including the extant aye-aye, Daubentonia as first reported here. So it comes as no surprise when Luo et al. report, “Eleutherodontids show a marked similarity to the primate Daubentonia in the ventrally bent rostrum and deep mandible, and both features are interpreted to be reinforcement for incisor gnawing.” That’s the case only with Vilevolodon this time. Others may be by convergence.

Molars
The jaw joint of the rodent allows for rostral-caudal and dorsal-ventral motion of the jaws. Luo et al. report, in Villevolodon it is not possible for the mandible to move posteriorly or horizontally, but their images show a continuous anteroposterior trough/furrow in the three molars, though not to the extent seen in sister taxon Shenshou. Molars with a long and continuous trough for rostral-caudal grinding appear by convergence in several reptile/mammal clades.

Incisor replacement
Luo et al. report, “Incisor replacement is prolonged until well after molars are fully erupted, a timing pattern unique to most other mammaliaforms. In rodents incisors never stop growing. The growth pattern in Vilevolodon may be the first step toward that. Not sure why Luo et al. are missing all these strong rodent clues.

Gliding?
Meng et al. 2017 note: “They [Vilevolodon and kin] are the most primitive known gliders in mammal evolution, evolving approximately 100 million years before the earliest known therian gliders.” Earlier, with the appearance of the stem pangolin, Zhangheotherium at the start of the Cretaceous, the ghost lineage for primates, flying lemurs and bats was also set to that time or earlier. Before the advent of flying birds, but after the advent of predatory theropods, many mammals had evidently taken to the trees. And one way to get from tree to tree without descending to the dangerous turf is to jump, glide and fly. I predict we’ll find the big-handed ancestors of bats in Jurassic and Cretaceous strata someday. They are already volant shortly after the K-T extinction event.

Hearing in Vilevolodon
With the reappearance of post-dentary bones in taxa like Vilevolodon, the auditory acuity that was more highly developed in its ancestors must have suffered a setback. By the evidence provided, the massive jaw joint must have been more important for its survival.

Figure 8. Multituberculate Kryobaatar mandible in lateral and medial views. Here post-dentary bones are absent. The malleus (quadrate) and ectotympanic are on the skull.

Figure 4. Multituberculate Kryobaatar mandible in lateral and medial views. Here post-dentary bones are absent here. The malleus (quadrate) and ectotympanic are on the skull.

Getting back to the purported patagium of Maiopatagium
which we looked at yesterday. It is not apparent and the authors do not describe it. Rather, Meng et al. 2017 sidestep this by reporting, “Furthermore, we report a second eleutherodont specimen (BMNH2942) preserved with a halo of carbonized fur and patagial membranes, similar to those of Maiopatagium.” The patagial taxon remains unnamed in the Maiopatagium paper (Meng et al. 2017), but is named in a second paper appearing on the same day. It is today’s subject, Vilevolodon (Fig. 1)

References
Filan SL 1991. Development of the middle ear region in Monodelphis domestica (Marsupialia, Didelphidae): marsupial solutions to an early birth. Journal of Zoology 225(4): 577–588 DOI: 10.1111/j.1469-7998.1991.tb04326.x
Luo Z-X, Meng Q-J, Grossnickle DM, Neander AI, Zhang Y-G and Ji Q 2017. New evidence for mammaliaform ear evolution and feeding adaptation in a Jurassic ecosystem. doi:101.1038/nature 23483\
Meng Q-J, Grossnickle DM, Liu D, Zhang Y-G, Neander AI, Ji Q and Luo Z-X 2017.
New gliding mammaliaforms from the Jurassic. Nature (advance online publication)
doi:10.1038/nature23476
Jenkins FA, Jr, Gatesy SM, Shubin NH and Amaral WW 1997. Haramiyids and Triassic mammalian evolution. Nature 385(6618):715–718.
Luo Z-X, Gatesy SM, Jenkins FA, Jr, Amaralc WW and Shubin NH 2015. Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution. PNAS 112 (51) E7101–E7109.

wiki/Haramiyavia
wiki/Vilevolodon
wiki/Maiopatagium

Figuring out the upside-down skull of Yanoconodon

Figure 1. Yanoconodon fossil in situ. See the skull in closeup in figure 2.

Figure 1. Yanoconodon fossil in situ. See the skull in closeup in figure 2. The published tracing is distorted here to match the underlying photo.

Wikipedia reports, “Yanoconodon was a small mammal, barely 5 inches (13 centimetres) long. It had a sprawling posture, Yanoconodon was a Eutriconodont, a group composing most taxa once classified as “triconodonts” which lived during the time of the dinosaurs. These were a highly ecologically diverse group, including large sized taxa such as Repenomamus that were able to eat small dinosaurs, the arboreal Jeholodens, the aerial volaticotherines and the spined Spinolestes. Yanoconodon is inferred to be a generalized terrestrial mammal, capable of multiple forms of locomotion.

Figure 1. Yanoconodon is exposed in ventral view. Even so, if you employ DGS, even on a fuzzy photo, you can put together a reconstruction that shares several traits with Repenomamus.

Figure 2. Yanoconodon is exposed in ventral view. Even so, if you employ DGS, even on a fuzzy photo, you can put together a reconstruction that shares several traits with Repenomamus.

Mammal-like reptiles?
Wikipedia also reports, “The Yanoconodon holotype is so well preserved that scientists were able to examine tiny bones of the middle ear. These are of particular interest because of their “transitional” state: Yanoconodon has fundamentally modern middle ear bones, but these are still attached to the jaw by an ossified Meckel’s cartilage. This is a feature retained from earlier stem mammals, and illustrates the transition from a basal tetrapod jaw and ear, to a mammalian one in which the middle ear bones are fully separate from the jaw. Despite this feature Yanoconodon is a true mammal. It is thought that the feature was retained during early embryo development,[4] whereas it is lost in most other mammal groups. The intermediate anatomy of the middle ear of Yanocodon is said to be a “Rosetta Stone”[5] of mammalian middle ear evolution.”

In the large reptile tree (LRT, 1037 taxa) Yanoconodon, Repenomamus, Jeholodens and Spinolestes are not mammals, but very close to the base of the Mammalia. Both clades share Pachygenelus as last common ancestor. So that means the ‘transitional state’ mentioned above is indeed outside the Mammalia. Other paleontologists consider this list of taxa to be mammals, but here the mammal-like traits they had were developed in parallel and not quite to mammal standards.

Figure 4. Repenomamus reconstructed using DGS methods. The manus and feet are loose figments at present. Despite its predatory nature, note the reduction in canines, a clade trait.

Figure 4. Repenomamus reconstructed using DGS methods. The manus and feet are loose figments at present. Despite its predatory nature, note the reduction in canines, a clade trait.

The skull of Yanoconodon
(Fig. 2) can be largely, but not completely, reconstructed based on the visible bones. The skull is low and wide and without the typical constriction anterior to the jugals. The anterior teeth are large and spike-like while the posterior teeth are molariform. Large teeth typically require deep roots and deep bones to house those roots. The mandibles are as long as the skull. The small orbits are far forward on the skull and the temporal fenestra are correspondingly large.

Figure 2. The origin and radiation of stem mammals and crown mammals. Compare the LRT tree (above) to a recent cladogram by Close et al. 2015.

Figure 2. The origin and radiation of stem mammals and crown mammals. Compare the LRT tree (above) to a recent cladogram by Close et al. 2015.

With the new data on Yanocondon
several taxa within the LRT shifted places, but not far and still within the derived Cynodontia. Something about the Mammalia helped them survive several extinction events that the derived Tritylodontia (= Pseudomammalia) succumbed to. Pseudomammalia LOOK like mammals, but are not mammals. They continued to exist into the Early Cretaceous and some, like Repenomamus, were quite large.

References
Close RA, Friedman M, Lloyd GT and Benson RBJ 2015. Evidence for a mid-Jurassic adaptive radiation in mammals. Current Biology. 25(16): 2137–2142. 
Luo Z, Chen P, Li G, and Chen M 2007.
 A new eutriconodont mammal and evolutionary development in early mammals. Nature 446:15. online Nature

wiki/Yanoconodon

Vintana and the vain search for the clades Allotheria and Gondwanatheria

Figure 1. Vintana as originally illustrated. I added colors to certain bones. Note the high angle of the ventral maxilla and the deep premaxilla. Lateral view reduced to scale with other views.

Figure 1. Vintana as originally illustrated. I added colors to certain bones. Note the high angle of the ventral maxilla and the deep premaxilla. Lateral view reduced to scale with other views.

Earlier we looked at Vintana (Fig. 1, Krause et al. 2014a, b). To Krause et al. Vintana represented the first specimen in the clades Allotheria and Gondwanatheria to be known from more than teeth and minimal skull material.

To Krause et al. 
Allotheria included Multituberculata and nested between the clade Eutriconodonta (including Repenomamus and Jeholodens) and the clade Trechnotheria (including the spalacotheres Maotherium and Akidolestes) and Cronopio, Henkelotherium, Juramaia, Eomaia, Eutheria and Metatheria.

Taxon exclusion issues
The large reptile tree (LRT, 1005 taxa) did not recover the above clades or relationships. Alotheria does not appear in the LRT.

  1. Multituberculata, Henkelotherium and Maotherium nest within Glires (rats and rabbits and kin) in the LRT.
  2. Repenomamus and Jeholodens nest within the pre-mammalian trityllodontid cynodonts in the LRT.
  3. Akidolestes nests within basal Mammalia, close to Ornithorhynchus in the LRT.
  4. Cronopio and Juramaia nest within basal Mammalia between Megazostrodon and Didelphis in the LRT.
  5. Eomaia nests at the base of the Metatheria in the LRT.
  6. Vintana nests with Interatherium among the derived Metatheria (marsupials), with wombats, like Vombatus and Toxodon in the LRT.

Despite a paper in Nature
and a memoir of 222 pages in the Journal of Vertebrate Paleontology; despite CT scans and firsthand examination with electron microscopes; despite being examined and described by many of the biggest name and heavy hitters in paleontology… Krause et al. never understood that Vintana was just a derived wombat, evidently due to taxon exclusion problems.

Figure 3. Interatherium does not nest with notoungulates or other purported interotheres. Rather cat-sized Interatherium nests with wombats, between Vombatus and the giant Toxodon.

Figure 2. Interatherium does not nest with notoungulates or other purported interotheres. Rather cat-sized Interatherium nests with wombats,with Vintana,  between Vombatus and the giant Toxodon

The large reptile tree now includes
1005 taxa, all candidates for sisterhood with every added taxon. Despite the large gamut of 74 taxa employed by Krause et al. they did not include the best candidates for Vintana sisterhood. Perhaps the fault lies in the reliance of prior studies and paradigms. Perhaps the fault lies in the over reliance by Krause et al. and other mammal workers, on dental traits. Perhaps the fault lies in the absence of pertinent sisters to the above-named taxa, including Interatheriium for Vintana.

In any case
Vintana does not stand alone as the only taxon in its clade represented by skull material. Based on its sisterhood with Interatherium, we have  pretty good idea what its mandibles and post-crania looked like. Yes, Vintana is weird. But Interatherium is also weird in the same way, just not as weird.

The LRT has dismantled and invalidated
several other clades, too, Ornithodira and Parareptilia among them.

References
Krause DW, Hoffmann S, Wible JR, Kirk EC, and several other authors 2014a. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature. online. doi:10.1038/nature13922. ISSN 1476-4687.
Krause DW et al. 2014b. Vintana sertichi (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology Memoir 14. 222pp.

wiki/Vintana
pterosaur heresies – Vintana

Danjiangia: not a chalicothere, not a brontothere…

It’s an early Eocene pig
according to the large reptile tree (LRT, 1003 taxa). A large gamut minimizes inclusion set bias and gains greater authority with every added taxon. It also reduces the average phylogenetic distance between taxa, all of which are species and individuals, not suprageneric taxa.

Figure 1. Danjiangia nests with the extant pig, Sus, in the LRT.

Figure 1. Danjiangia nests with the extant pig, Sus, in the LRT. Note the very low naris and nasal. The lost skull could have been elevated, as imagined here after phylogenetic analysis.

Danjiangia pingi (Wang 1995; early Eocene) was originally described as a basal chalicothere with brontothere traits. Then Hooker and Dashzeveg (2003) nested it as a basal brontothere (without including any other brontotheres). Mihlbacher 2008 and others used it as an outgroup to the brontotheres. The posterior skull is not known, but note the rise over the orbits suggesting a tall cranium, as in Sus (Fig. 2). Also note the very low naris below the low nasals. Usually you don’t see nasals so low, and perhaps that is due to taphonomic shifting.

Figure 2. Skull of the extant pig, Sus in several views.

Figure 2. Skull of the extant pig, Sus in several views. Note the elevated cranium and squamosal.

The long fused dentary 
of Danjiangia is a trait also shared with pigs and other taxa, like chalicotheres, by convergence.

Figure 1. Skeleton of Sus, the pig, a taxon commonly used as an outgroup for whales. In the LRT it is a sister to other even-toed ungulates, like Giraffa, not Odontoceti nor Mysticeti.

Figure 3. Skeleton of Sus, the pig. It provides good clues as to the missing postcranial skeleton of Danjiangia.

Sus the pig
(Fig. 3) provides good clues as to the missing postcranial skeleton of its sister taxon, Danjiangia. The other model for post-cranial details is the basal artiodactyl, Cainotherium (Fig. 4).

Fig. 1. Cainotherium nests at the base of the Artiodactyla or even-toed ungulates. I wonder if it had five fingers, even if vestiges, given that Ancodus, a derived artiodactyl, retains five fingers.

Fig. 4. Cainotherium nests at the base of the Artiodactyla or even-toed ungulates. I wonder if it had five fingers, even if vestiges, given that Ancodus, a derived artiodactyl, retains five fingers.

Why was the pig connection missed by others?
For the same reason that modern workers continue to include pterosaurs with archosaurs. It’s a tradition. Nobody wants to do the extra work of testing other candidate taxa. Nobody wants to acknowledge contrarian studies. Paleontology tends to run very slowly as we learned earlier here. Hail, hail the status quo!

Figure 1. Subset of the LRT focusing on ungulates, which split into three clades here.

Figure 5. Subset of the LRT focusing on ungulates, which split into three clades here. Note the nesting of Sus together with Danjiangia.

References
Beard KC 1998. East of Eden: Asia as an important center of taxonomic origination in mammalian evolution; pp. 5–39 in Beard and Dawson (eds.), Dawn of the Age of Mammals in Asia. Bulletin of Carnegie Museum of Natural History 34.
Mihlbachler MC 2004. Phylogenetic systematics of the Brontotheriidae (Mammalia, Perissodactyla). PhD dissertation. Columbia University. p. 757.
Mihlbachler MC 2008. Species taxonomy, phylogeny and biogeography of teh Brontotheriidae (Mammalia: Perissodactyla). Bulletin of the American Museum of Natural History 311:475pp.
Wang Y 1995. A new primitive chalicothere (Perissodactyla, Mammalia) from the early Eocene of Hubei, China. Vertebrata Palasiatica 33: 138–159.

Stylinodon may be a giant herbivorous mink

In 1873 
O. C. Marsh 1874) found an extinct Eocene (50.3 to 40.4 Ma) mammal “of great interest. The lower molar teeth, all essentially alike, and inserted in deep sockets” were the most striking feature. He named it Stylodon mirus (Figs. 1,2). All the teeth grew with “persistent pulps” and had a thin layer of enamel. The specimen was considered close to Toxodon with some edentate affinities (Marsh 1897). Stylinodon was placed under the family Stylinodontidae and the order Tillodontia. According to Schoch 1986 (first issue of JVP!) its ancestors were like Onychodectes.

Stylinodon mirus (Marsh 1874; middle Eocene, 45 mya; Figs. 1-2) was originally considered a taeniodont, perhaps derived from Onychodectes. Here it nests with Mustela, the living European mink, among the Carnivora. There were twice as many molars (4), each with a single root, as in the two double rooted molars of the mink. Large claws and certain forelimb traits indicate that Stylinodon was a digger, not a cursor.

The present nesting
of Stylinodon mirus (YPM VP 011095, Marsh 1874; Figs. 1, 2) in the Carnivora occurred when I realized it was a poor fit at the base of the Condylarthra/Paenungulata, despite its herbivorous dentition and tusk-like teeth (canines, not incisors).

Figure 1. Stylinodon skull. Note the transverse premaxilla, a trait of the Carnivora.

Figure 1. Stylinodon skull. Note the transverse premaxilla, a trait of the Carnivora.

Distinct from condylarths
Stylinodon has a transverse premaxilla, essentially invisible in lateral view. The lower canine is the anteriormost tooth on the dentary. These traits are shared with other members of the Carnivora. In the present taxon list Stylinodon shares more traits with Mustela, the European mink (Fig. 1) despite the loss of molar cusps and increase in size. They both were diggers. Together they nest with Phoca, the seal, and Palaeosinopa, the amphibious piscivore, all derived from a sister to Procyon, the omnivorous raccoon (Fig. 2).

Figure 1. Stylinodon compared to Mustela, the European mink to scale.

Figure 2 Stylinodon compared to Mustela, the European mink to scale.

As in the earlier issue
with indricotheres, related taxa can have distinctively different types of teeth, one more reason to not weight dental traits too heavily, unless that’s all you have.

Figure 2. Mustela the European mink is an extant relative to Stylinodon.

Figure 3. Mustela the European mink is an extant relative to Stylinodon.

Mustela lutreola (Linneaus 1761; extant European mink; up to 43cm in length) is a fast and agile animal related to weasels and polecats. Mustela lives in a burrow, but it also swims and dives skilfully. It is able to run along stream beds, and stay underwater for one to two minutes. Mustela is derived from a sister to Phoca and other seals, all derived from a sister to Procyon. With this close relationship, Stylinodon (Fig. 2 a giant weasel with simple teeth.

Schoch and Lucas 1981
and Schoch 1983 considered Stylinodon and kin derived from a sister to the long-legged basal condylarth, Onychodectes. The large reptile tree (LRT, Fig. 2) does not support that nesting. Onychodectes has a long premaxilla lacking in taeniodonts.

Figure 2. Subset of the LRT showing the Carnivora nesting at the base of the Eutheria (placental mammals).

Figure 4. Subset of the LRT showing the Carnivora nesting at the base of the Eutheria (placental mammals).

Schoch and Lucas 1981
determined that Stylinodon had two upper incisors (one lower), a giant canine, four premolars and three molars, as in Onychodectes. That may be so, but the premolars and molars look alike.

 

Figure 6. Wortmania as drawn freehand by Schoch compared to bones Photoshopped together.

Figure 6. Wortmania as drawn freehand by Schoch compared to bones Photoshopped together.

Wortmania (Hay 1899, Williamson and Brusatte 2013; above) and Psittacotherium (Cope 1862; below) are related to Stylinodon. All are among the largest taxa in the early post-Cretaceous, derived from smaller weael-like basal mammals in the Cretaceous.

Figure 7. Psittacotherium in various views.

Figure 6.  Psittacotherium in various views. Overall it is elongated to more closely match related taxa.

It is rare but not unheard of
for members of the Carnivora to become omnivores and herbivores. Think of the giant panda and certain viverrids. Now the stylinodontid taeniodonts join their ranks.

References
Linneaus C von 1761. xxx
Marsh OC 1874. Notice of new Tertiary mammals 3. American Journal of Science. (3) 7i: 531-534.|
Marsh OC 1897. The Stylinodontia, a suborder of Eocene Edentates. The American Journal of Science Series 4 Vol. 3:137-146.
Rook DL and Hunter JP 2013. Rooting Around the Eutherian Family Tree: the Origin and Relations of the Taeniodonta. Journal of Mammalian Evolution: 1–17.
Schoch RM and Lucas SG 1981. The systematics of Stylinodon, an Eocene Taeniodont (Mammalia) from western North America. Journal of Vertebrate Paleontology 1(2):175-183.
Schoch RM 1983. Systematics, functional morphology and macroevolution of the extinct mammalian order Taeniodonta. Peabody Museum of Natural History Bulletin 42: 307pp. 60 figs. 65 pls.

 

 

 

wiki/Stylinodon
wiki/Mustela