Masrasector: not a placental… a killer marsupial

Of course, by that I mean,
skunk-sized Masrasector nananubis (Fig. 1; Simons and Gingerich 1974; Borths and Seiffert 2017), like other hyaenodonts in the LRT (Fig. 2), is also a marsupial… a descendant of a sister to the quiet little, nocturnal Virginia opossum (Didelphis, Fig. 2), but closer to Borhyaena.

[Note: everyone agrees that hyaenodonts are NOT related to hyaenas.]

Taxon exclusion,
based on tradition, evidently keeps marsupials out of current phylogenetic analyses of hyaenodonts. Hopefully that will be corrected, or at least tested, in the future.

Hyaenodonts,
like Masrasector, are traditionally considered placental (eutherian) carnivorous creodonts. They have not one, but three carnassial teeth.

According to the LRT,
that was by convergence. Certain bats also have carnassial teeth. So it can happen.

By contrast
the large reptile tree (LRT, 1066) nests Masrasector and all other hyaenodonts as marsupial carnivores. Borths and Seiffert (2017) did not expand their taxon list to include marsupials and thus, taxon exclusion implanted a deadly flaw in their otherwise brilliant and technical paper.

Figure 3. Arctocyon is no longer an ungulate placental, but a carnivorous marsupial, close to Thylacinus.

Figure 2. Arctocyon is no longer an ungulate placental, but a carnivorous marsupial.

Borths and Seiffert note:
“Body mass in hyaenodonts is difficult to estimate because there are no living taxa analogous to these large-headed placental carnivores with multiple carnassials.”

“The most surprising part of this study was that Masrasector and its kin are part of a massive radiation of hyaenodonts that originated in Africa.” 

“Hyaenodonts have been a little neglected as a group.”

Wikipedia reports on the history of the Creodonta:
“Creodonta” was coined by Edward Drinker Cope in 1875. Cope included the oxyaenids and the viverravid Didymictis but omitted the hyaenodontids. In 1880. he expanded the term to include MiacidaeArctocyonidaeLeptictidae (now Pseudorhyncocyonidae), OxyaenidaeAmbloctonidae and Mesonychidae.[12] Cope originally placed creodonts within the Insectivora. In 1884, however, he regarded them as a basal group from which both carnivorans and insectivorans arose. Hyaenodontidae was not included among the creodonts until 1909. Over time, various groups were removed, and by 1969 it contained, as it does today, only the oxyaenids and the hyaenodontids.”

Figure 2. Masrasector nests with Borhyaena in the marsupial clade.

Figure 3. Masrasector nests with Borhyaena in the marsupial clade.

Earlier we looked at members at the base of this clade, like Amphicyon and Arctocyon (Fig. 2), giant closer descendants of Didelphis. Hyaenodonts and other creodonts are different from members of the Carnivora because they’re not placentals, something that has escaped the notice of paleontologists. Creodonts / hyaenodonts appear earlier in the fossil record because marsupials appear before placentals.

References
Borths MR, Seiffert ER 2017. Craniodental and humeral morphology of a new species of Masrasector (Teratodontinae, Hyaenodonta, Placentalia) from the late Eocene of Egypt and locomotor diversity in hyaenodonts. PLoS ONE 12(4): e0173527.
Simons EL, Gingerich PD 1974. New carnivorous mammals from the Oligocene of Egypt. Annals of the Geological Survey of Egypt. 1974; 4: 157–166.
Author interview online here.
Pasttime.org podcast interview here.

Juehuaornis traced and reconstructed from lo-rez data

Figure 1. Juehuaornis in situ with tracing

Figure 1. Juehuaornis in situ with tracing

Juehuaornis zhangi (Wang et al. 2015) is a new ornithuromorph genus from the Early Cretaceous of western Liaoning, China.

Figure 2. Juehuaornis reconstructed. Note the scale bars. This is a tiny bird.

Figure 2. Juehuaornis reconstructed. Note the scale bars. This is a tiny bird.

From the abstract: Here we report on a new basal ornithuromorph bird, Juehuaornis zhangi gen et sp. nov.,based on a nearly complete and articulated subadult skeleton from the Lower Cretaceous Jiufotang Formation in Lingyuan of western Liaoning, China. It displays ornithomorph synapomorphies, such as a synsacrum composed of 12 sacral vertebrae, a short pygostyle , long and curved scapula, U-shaped furcula without a hypocleidum, coracoid with a developed procoracoid process and a concaved lateral margin,a keel extended along the full length of sternum, major and minor metacarpals fused proximally and distally, and proximal phalanx of digit II expanded caudally. The new specimen is distinguishable from other known ornithuromorphs by some unique features including a long rostrum for approximately 70% the total length of the skull, cranial end of upper jaw hooked, and cranial end of lower jaw straight. The new specimen provides new important morphological information regarding Ornithomorpha, and it represents a new ecotype of this group.

Figure 3. This is a very low rez image of the skull, the best I could wrangle from the original paper.

Figure 3. This is a very low rez image of the skull, the best I could wrangle from the original paper. That’s a hand over the skull. See figure 1. Not sure about the described premaxilla tip described in the paper. Requested high-rez data will replace this if it comes.

In the large reptile tree,
Juehuaornis nests with other small Early Cretaceous tobirds between Longicrusavis and higher toothed birds like Changzuiornis and Yanornis.

References
Wang R-F, Wang Y and Hu Dong-yu 2015. Discovery of a new ornithuromorph genus, Juehuaornis gen. nov. from Lower Cretaceous of western Liaoning, China. Global Geology 34(1):

Eudimorphodon skull reconstructed

Figure 1. Eudimorphodon ranzii nests at the base of all non-dimorphodontid pterosaurs. Here bones are colorized using DGS and reconstructed below in several views. I can't identify all the bones here, just the easy ones. Red spot in orbit is the rotated pterygoid with teeth. Other pterosaurs don't have such teeth, but then other pterosaurs don't have such marginal teeth either. Line art from F. Dalla Vecchia.

Figure 1. Eudimorphodon ranzii nests at the base of all non-dimorphodontid pterosaurs. Here bones are colorized using DGS and reconstructed below in several views. I can’t identify all the bones here, just the easy ones. Red spot in orbit is the rotated pterygoid with teeth. Other pterosaurs don’t have such teeth, but then other pterosaurs don’t have such marginal teeth either. Line art from F. Dalla Vecchia.

This is Eudimorphodon (Late Triassic), the basalmost of all the non-dimorphodontid pterosaurs. The in situ skull is crushed (Fig. 1), so if you want to see what it looks like in three views you take the parts and put the model back together.

Figure 4. Lateral, dorsal and cross-sectional views of Eudimorphodon ranzii. Note the overlap of the posterior ribs over the hind limbs and the very wide torso. The cross section shows the 2nd dorsal ribs and the 23rd. Note the small ischium which could only produce small eggs. A little taller and wider than we thought before. The forelimbs are pretty short relative to the torso.

Figure 2. Lateral, dorsal and cross-sectional views of Eudimorphodon ranzii. Note the overlap of the posterior ribs over the hind limbs and the very wide torso. The cross section shows the 2nd dorsal ribs and the 23rd. Note the small ischium which could only produce small eggs. A little taller and wider than we thought before. The forelimbs are pretty short relative to the torso.

Eudimorphodon-pterygoid-teeth588.jpg

Close up of Eudimorphodon jugal and pterygoid (yellow) from an old photocopy from back in the day. 

We looked at
the post-cranium of Eudimorphodon earlier here. The torso is weirdly flattened, like that of Sharovipteryx, making the entire body a wing shape.

Figure 3. Sharovipteryx reconstructed. Note the flattened torso.

Figure 3. Sharovipteryx reconstructed. Note the flattened torso.

References
Wild R 1978. Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen Trias von Cene bei Bergamo, Italien. Bolletino della Societa Paleontologica Italiana 17(2): 176–256.
Zambelli R 1973. Eudimorphodon ranzii gen.nov., sp.nov. Uno Pterosauro Triassico. Rendiconti Instituto Lombardo Accademia, (rend. sc.) 107: 27-32.

wiki/Eudimorphodon

 

You heard it here first: No two Archaeopteryx look the same.

The science section
of the online British news outlet, the Guardian, reported on the 12th (Haarlem) specimen of Archaeopteryx re-named Ostromia. You can read that story online here.

From the Guardian article:
“Of the Twelve Specimens Once Known as Archaeopteryx (SPOKA), only nine continue to carry that name. At the end of the day, we can’t all be winners. But even within that group of nine specimens, no two Archaeopteryx look the same. Rauhut and colleagues report that there is significant variation in the size, shape, spacing and orientation of the teeth, as well as differences in body size between the different specimens. This could be an ontogenetic pattern, with larger individuals representing adults with more developed dentition. Alternatively, as the Solnhofen Basin constituted a tropical island archipelago during the Late Jurassic, these differences in body size and dentition could be interpreted as island adaptations. Similarly to today’s Galápagos finches, different populations of Archaeopteryx may have adapted to different insular environments.”

We looked at
Solnhofen birds (Fig. 1) earlier here and Ostromia here. Since 2015 readers have known that no two Archaeopteryx specimens were identical and that phylogenetic analysis split them apart to nest at the base of each one of all the Cretaceous bird clades. And yes, we know of an embryo archaeopterygid, the Liaoning embryo most closely related to the London specimen.

Figure 3. Several Solnhofen birds, including Archaeopteryx, compared to Ostromia to scale.

Figure 3. Several Solnhofen birds, including Archaeopteryx, compared to Ostromia to scale.

It really is time to
run these birds through analysis and either affirm, modify or invalidate the results of the large reptile tree. And it should be done by someone with firsthand access to all the specimens. That would be a good test.

References
Elzanowski, A., 2002. Archaeopterygidae (Upper Jurassic of Germany) In: Chiappe LM, Witmer LM, eds. Mesozoic Birds. Above the Heads of Dinosaurs. Berkeley: University of California Press. 129-159.
Foth C, Rauhut OWM. 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology 17:236

https://www.theguardian.com/science/2018/feb/21/the-new-specimen-forcing-a-radical-rethink-of-archaeopteryx

Redefining what makes a dinosaur

Ran across this online article (citation below)
summarized: “The once-lengthy list of “definitely a dinosaur” features had already been dwindling over the past few decades thanks to new discoveries of close dino relatives such as Teleocrater. With an April 2017 report of Teleocrater’s skull depression (SN Online: 4/17/17), yet another feature was knocked off the list.”

Evidently the only trait that is still on the list is a perforated acetabulum.

My résponse:
long time readers will recognize this answer:

The dear departed Dr. Larry Martin used to play this game. He’d say ‘tell me a character you think defines a clade and I’ll give you an exception.’ In dinosaur pelves, ankylosaurs are the exception that do not have a perforated acetabulum. The lesson: You can’t define a clade by a single or a dozen character traits. 

You can define a clade using a cladogram. A cladogram uses hundreds of traits to recover relationships including “the last common ancestors and all of its descendants.” On that basis Dinosauria include Herrerasaurus at the base and the first dichotomy splits Theropoda from Phytodinosauria. The proximal outgroup is the Crocodylomorpha, basal members of which were small and bipedal, like early dinosaurs. That means Archosauria includes only dinos and crocs. Teleocrater is in the lineage of stem Archosauria. Unfortunately, prior workers excluded many relevant taxa, which is why they did not recover these relationships. Cladogram, links and more data here: 
The last few items on the dinosaur list:
  1. Until Teleocrater came along, only dinosaurs were known to have a deep depression at the top of the skull, an attachment site for some jaw muscles probably related to bite strength.
  2.  Dinosaurs and some other dinosauromorphs such as Silesaurus opolensis have an enlarged crest on the upper arm bone where muscles attached
  3. Along with dinosaurs, dinosauromorphs S. opolensis and Asilisaurus kongwe may have had epipophyses, bony projections at the back of the neck vertebrae.
  4. An extra (fourth) muscle attachment site, called a trochanter, at the point on the femur that meets the hip is also found in dinosauromorph Marasuchus lilloensis.

Sources: S.J. Nesbitt et al/Nature 2017; S.L. Brusatte et al/Earth-Science Reviews 2010

Taxon exclusion. Phylogenetic analysis. Yada-yada. 

References
https://www.sciencenews.org/article/new-fossils-are-redefining-what-makes-dinosaur

Nesbitt et al. 2017 The earliest bird-line archosaurs and the assemblof the dinosaur body plan. Nature (Teleocrater paper).

When turtles lost their teeth

When animals lose something,
be it a tail, finger, limbs, eyes or teeth, usually a vestige is left behind.

When turtles lost their ancestral teeth,
they should have left empty alveoli along their jaw rims. And the place to look for empty alveoli in turtles is in the most primitive turtle in the large reptile tree, the late-surviving Niolamia (Fig. 1), one of the great horned meiolaniid turtles.

Figure 1. Palate of the basal turtle Niolamia with arrows pointing to pinprick alveoli lacking teeth.

Figure 1. Palate of the basal turtle Niolamia with arrows pointing to pinprick alveoli lacking teeth.

Tiny pinpricks
along the maxilla (Fig. 1) seem to show where tiny teeth once erupted in Niolamia.

Earlier we looked at similar alveoli in the jaw tips of a gray whale where desmostylian tusks once emerged.

Helpless and able newborn mammals

I’m going to crowd source this one,
but I think I covered all the bases here. In this subset of the large reptile tree (LRT, 1165 taxa) I’ve divided placental mammals born helpless (blue) from mammals born able to walk, swim and see (pink). I’ll need your help if there are any exceptions, like pangolins, that I missed one way or the other. Fossils are colorized based on phylogenetic bracketing.

Figure 1. Newborn mammals are born either helpless, like humans, or able to keep up with their mother, like horses. I think I located the split correctly here. Let me know I missed a few.

Figure 1. Newborn mammals are born either helpless, like humans, or able to keep up with their mother, like horses. I think I located the split correctly here. Let me know I missed a few. Fossil taxa are colored based on phylogenetic bracketing. 

Marine taxa need to be ready to go from the first minute.
Apparently so do the large plant-eaters ( including ant and copepod eaters), beginning with long-legged former tree shrew, Onychodectes.

Dens and nests
are associated with basal mammals, like us. Not so much with the derived herbivores (and anteaters) of the plains and forests. All of them get milk from their mothers before they start to dine on meat, plants, ants and copepods. Some of them have to keep up with here. Some of them have to keep up with her underwater.

BTW
there also seems to be a behavioral node at Maelestes in which succeeding taxa are all leaving the trees for good. Of course, that also happens exceptionally with the various mole and aquatic clades in more basal mammals.

Where are the auditory bullae in desmostylians?

A reader wondered about
tympanic (auditory) bullae (ear container bones) in desmostylians. Then I wondered, too. All whales are famous for having them. Nobody talks about them in desmostylans. So what gives? Here are the data:

Short answer:
apparently bullae are easily knocked off and/or ignored during the process of fossilization and extraction, both in mysticetes and desmostylians. Some examples follow:

Gray whale (Eschrichtius)
Bullae were present, but somehow got knocked off when it came time to draw the diagram (Fig. 1).

Figure 1. Gray whale (genus: Eschrichtius) in which bullae were present, but omitted from a palate diagram.

Figure 1. Gray whale (genus: Eschrichtius) in which bullae were present, but omitted from a palate diagram.

Cornwallius (a pre-desmostylian cambaythere) — overlooked bulla, called a ‘mass’ in the text.

Figure 3. The pre-desmostylian Cornwallius. Here the tympanic bulla (bright green) was considered "a mass" in the text and otherwise was not labeled.

Figure 3. The pre-desmostylian Cornwallius. Here the tympanic bulla (bright green) was considered “a mass” in the text and otherwise not labeled.

Neoparadoxia (basal desmostylian)
Here you can see the depression that receives the bullae, but the bullae became missing at some stage in the process.

Figure 3. Palate of Neoparadoxia, a basal desmostylian, apparently missing the tympanic bullae (ear bones).

Figure 3. Palate of Neoparadoxia, a basal desmostylian, apparently missing the tympanic bullae (ear bones). Note the ear canal bones extending laterally, as in the hippo (figure 6).

Desmostylus, a derived desmostylian close to right whales
Same here. Bulla not published. Depression for the reception still present.

Figure 4. Desmostylus with missing bullae replaced in the empty spots left behind.

Figure 4. Desmostylus with missing bullae replaced in the empty spots left behind. Skull is obviously distorted and missing a big part of the cranium.

Caperea, a basal right whale
Here’s an odd one. Not sure what happened to the bulla in ventral view. They seem to appear in occiput view.

Figure 6. Caperea, a basal right whale, apparently missing the bullae in palate view that it had in occipital view.

Figure 5. Caperea, a basal right whale, apparently missing the bullae in palate view that it had in occipital view. If not, please advise.

Hippopotamus
This goes back somewhat on the tree, but hippos are in the lineage of baleen whales in the LRT and their auditory bones are present.

Figure 7. Hippopotamus with auditory meatus (ear canal) in green, bulla (ear bone container bones) in yellow.

Figure 6. Hippopotamus with auditory meatus (ear canal) in green, bulla (ear bone container bones) in yellow.

Ear bones compared
Baleen whale bullae greatly resemble toothed whale bullae. It’s true. Based on phylogeny, we’ll have to call this convergence. So is the loss of teeth in the rostrum of the sperm whale and blue whale. Convergence happens, but let’s keep an eye out for those bullae, now that we know what should be there.

Bipedal Cretaceous lizard tracks

These are the oldest lizard tracks in the world…
(if you don’t consider Rotodactylus (Early Triassic) strictly a ‘lizard’ (= squamate). One rotodactylid trackmaker, Cosesaurus, is a tiny lepidosaur).

Figure 1. Bipedal lizard tracks from South Korea in situ.

Figure 1. Bipedal lizard tracks from South Korea in situ. They are tiny.

From the abstract
“Four heteropod lizard trackways discovered in the Hasandong Formation (Aptian-early Albian), South Korea assigned to Sauripes hadongensis, n. ichnogen., n. ichnosp., which represents the oldest lizard tracks in the world. Most tracks are pes tracks that are very small. The pes tracks show “typical” lizard morphology as having curved digit imprints that progressively increase in length from digits I to IV, a smaller digit V that is separated from the other digits by a large interdigital angle. The manus track shows a different morphology from the pes. The predominant pes tracks, the long stride length of pes, narrow trackway width, digitigrade manus and pes prints, and anteriorly oriented long axis of the fourth pedal digit indicate that these trackways were made by lizards running bipedally, suggesting that bipedality was possible early in lizard evolution.”
Actually, the lizard was not running.
Typically in running tracks the prints are very far apart and these tracks are sometimes left toe to right heel.

Figure 2. Original and new tracings of the bipedal lizard tracks from South Korea. PILs are added,

Figure 2. Original and new tracings of the bipedal lizard tracks from South Korea. PILs are added. Manual digit 4 and 5 appear to have shifted.

 The authors did not venture who made the tracks.
They reported, “based on the palaeobiogeographic distribution of facultative extant families, the lizard that produced S. hadongensis tracks could well have been a member of an extinct family or stem members of Iguania, which was present in the Early Cretaceous.”
Actually the closest match among tested taxa
is with Eichstaettisaurus (Fig. 1), a basal member in the lineage of snakes. And this clade is close to the origin of geckos. ReptileEvolutiion.com and the large reptile tree would have been good resources for the authors to use. Lots of lizard pedes were illustrated and scored there.

Figure 3. Originally pictured as a generic lizard (below), here Eichstattsaurus scaled to the track size walks upright.

Figure 3. Originally imagined  as a generic lizard (below), here Eichstattsaurus matched and scaled to the track size walks upright.

 Based on a phylogenetic analysis of the tracks
the closest match in the LRT is with Eichstaettisaurus, so a slightly larger relative made them. Distinct from the skeletal taxon, the trackmaker had a longer p2.1 than 2.1 and pedal digit 1 was quite short. Otherwise a good match in all other regards.
So why walk bipedally?
It was walking, not running, so escape from predation can be ruled out. Elevating the upper torso and head, like a cobra, can be intimidating to rivals, or just offer a better view over local plant life. This sort of flexibility could have helped them get into the trees and then to move to higher branches.
References
Lee H-J, Lee Y-N, Fiorillo AR &  LÃ J-C 2018. Lizards ran bipedally 110 million years ago. Scientific Reports 8: 2617. doi:10.1038/s41598-018-20809-z

Cornwallius: not a desmostylian, an ancestor to desmostylians

These taxa
are part of the a recent review of mysticete (baleen whale) ancestors you can read about here, here and here.

Cornwallius sookensis (originally Desmostylus sookensis, Hay 1923, Cornwall 1922; Beatty 2006a, b; Early Oligocene, 25 mya; Fig. 1) was originally and traditionally considered a desmostylian (Fig. 3). Here it nests with Cambaytherium (Fig. 2), both basal to anthracobunids like Janjucetus. These taxa have a narrow skull and a deep jugal beneath the squamosal. The nares are anterior, rather than dorsal in location.

Figure 1. Adult Cornwallius look more like desmostylians. Juveniles look more like anthracobunids. Both are descendant taxa.

Figure 1. Adult Cornwallius look more like desmostylians. Juveniles look more like anthracobunids. Both are descendant taxa.

Note the resemblance
(lack of a downturned snout) on the juvenile to Cambaytherium (above). Apparently, neotony produces a straights-snout anthracobunid. Otherwise it evolves to the tusky, droop-snout, desmostylian grade.

Figure 2. Cambaytherium with a an alternate rostrum reversing taphonomic shifts.

Figure 2. Cambaytherium with a an alternate rostrum reversing apparent taphonomic shifts.

Beatty 2006
produced the following cladogram (Fig. 3) in which desmostylians are derived from the Moeritherium/Elephas clade. In the large reptile tree (LRT, 1163 taxa) cambaytheres and desmostylians arise from mesonychids and hippos.

Figure 2. From Beatty 2006b, a phylogeny of desmostylians derived from moeritherium, an aquatic relative of elephants and sirenians (manatees). Actually desmostylians arise from cambaytheres and anthracobunids, arising from hippos and mesonychids. 

Figure 3. From Beatty 2006b, a phylogeny of desmostylians derived from moeritherium, an aquatic relative of elephants and sirenians (manatees). Actually desmostylians arise from cambaytheres and anthracobunids, arising from hippos and mesonychids.

References
Beatty, BL 2006a. Rediscovered specimens of Cornwallius (Mammalia, Desmostylia) from Vancouver Island, British Columbia, Canada. Vertebrate Palaeontology. 1(1):1–6.
Beatty, BL 2006b. Specimens of Cornwallius sookensis (Desmostylia, Mammalia) from Unalaska Island, Alaska. Journal of Vertebrate Paleontology. 26(3):785–87.
Cooper LN, Seiffert ER, Clementz M, Madar SI, Bajpai S, Hussain ST, Thewissen JGM 2014. Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls. PLoS ONE. 9 (10): e109232. doi:10.1371/journal.pone.0109232. PMID 25295875.
Cornwall IE 1922. Notes on the Sooke Formation, Vancouver Island, B.C. Canadian Field Naturalist. 36:121–23.
Hay OP 1923. Characteristics of sundry fossil vertebrates. Pan-American Geologist. 39:101–120.
Kumar K 1991. Anthracobune aijiensis nov. sp. (Mammalia: Proboscidea) from the Subathu Formation, Eocene from NW Himalaya, India”. Geobios. 24 (2): 221–39. doi:10.1016/s0016-6995(91)80010-w. OCLC 4656806310.
Rose, KD et al. (8 other authors) 2014. Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India. Nature Communications. 5 (5570). doi:10.1038/ncomms6570.

wiki/Cambaytherium
wiki/Cornwallius