Fruitafossor: now a Late Jurassic echidna from Colorado

While reviewing the terrestrial descendants of tree shrews
yesterday, the Late Jurassic Fruitafossor (Figs. 1, 2) stuck out as a chronological misfit as it nested in the otherwise Tertiary edentates (= Xenarthrans).

Here is the problem,
and the solution.

A Jurassic edentate? No.
Fruitafossor windscheffeli (Luo and Wible 2005) used to nest in the LRT with digging edentates, like the armadillo-mimic, Peltephilus (Miocene), and for good reason…

Wikipedia reports,
“The teeth of Fruitafossor bear a striking resemblance to modern armadillos and aardvarks. Its vertebral column is also very similar to armadillos, sloths, and anteaters (order Xenarthra). It had extra points of contact among similar to the xenarthrous process that are only known in these modern forms.”

By contrast, Wikipedia concludes,
“Its shoulder-girdle is similar to a platypus or reptile, but many other features are more similar to most other modern mammals.”

What would Larry Martin say?
Run a complete analysis. Don’t rely on one, two or a dozen traits. And the Late Jurassic is so early in mammal evolution that it becomes important, too. There were fewer mammal clades back then. Edentates had not yet arrived.

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

Figure 1. Several drawings from Zhou and Wible that one must trust for accuracy. The verification data is too fuzzy to validate.

So is Fruitafossor a Late Jurassic edentate?
Or an edentate-mimic in the Late Jurassic?
With current scoring in the LRT, shifting Fruitafossor from the edentates to the base of the Monotremata adds 23 steps. Shifting to Early Cretaceous Lactodens within the Monotremata adds just 17 steps, the lowest number I could find. Lactodens has typical differentiated teeth and five fingers with small, sharp claws, traits not shared with Fruitafossor + edentates. Lactodens nests with the echidnas, Tachyglossus (extant, Figs. 3–5) and Cifelliodon (Early Cretaceous; Fig. 3). The latter has simple blunt teeth and the former is a known digger.

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.

So let’s reexamine scored traits… and solve this conundrum.
Has the LRT met its match? Very few skull traits are known from Fruitafossor. Even so, earlier I overlooked or mis-scored the following that gain importance in hindsight:

Fruitafossor:

  1. orbit contacts the maxilla
  2. 4 rather than 5 sacrals,
  3. coracoid present
  4. I could not score hind limb length without a pes and estimates won’t do
  5. proximal sesamoid of fibula present
  6. fibula diameter greater than half of tibia
  7. dorsal osteoderms absent (I misinterpreted scattered elements at Digimorph.org)

Tachyglossus:

  1. retroarticular process present as in Fruitafossor
  2. metacarpal 1 and 2 are the longest as in Fruitafossor
  3. longest manual digit 3 as in Fruitafossor
  4. manual digit 4 narrower than 3 as in Fruitafossor

Cifelliodon:

  1. three molars, as in Fruitafossor
Figure 1. Early Cretaceous Cifelliodon is ancestral to the living echidna, Tachyglossus according to the LRT. The lack of teeth here led to toothlessness in living echidnas. The skull of Tachyglossus is largely fused together, lacks teeth and retains only a tiny lateral temporal fenestra (because the jaws don't move much in this anteater. Compared to Cifelliodon the braincase is greatly expanded, the lateral arches are expanded and the two elements fuse, unlike most mammals.

Figure 3. Early Cretaceous Cifelliodon is ancestral to the living echidna, Tachyglossus according to the LRT. The lack of teeth here led to toothlessness in living echidnas. The skull of Tachyglossus is largely fused together, lacks teeth and retains only a tiny lateral temporal fenestra (because the jaws don’t move much in this anteater. Compared to Cifelliodon the braincase is greatly expanded, the lateral arches are expanded and the two elements fuse, unlike most mammals.

Figure 3. Tachyglossus skeleton, manus and x-rays. Note the perforated pelvis.

Figure 4. Tachyglossus skeleton, manus and x-rays. Note the perforated pelvis.

Figure 1. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws.

Figure 5. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws.

Results (as you might imagine, given these changes):
Fruitafossor is an edentate-mimic nesting basal to Cifellidon and Tachyglossus as a Late Jurassic echidna and monotreme in the LRT. Glad to get rid of that problem!

In their original description of Fruitafossor,
Luo and Wible 2005 nested their discovery between a monotreme clade and a clade with the mammal-mimic, Gobiconodon at its base, then a clade with another egg-laying mammal, Tinodon at its base, then a pangolin ancestor, Zhangheotherium, then a rabbit ancestor Henkelotherium, then two other monotremes, Dryolestes, Amphitherium and the carnivorous marsupial, Vincelestes.  Luo and Wible tested Tachyglossus, but not Cifelliodon, which was published in 2018. Note the simple, blunt teeth in Cifelliodon, nearly matching those in Fruitafossor. Given that the only fossil of Fruitafossor is a bit jumbled, it is possible that it, too, had five fingers in vivo, like other monotremes. With only four fingers (Fig. 1) Fruitafossor had a good excuse for pretending to be an edentate.

So, yes, the LRT was up to the challenge.
But it took insight, lacking until now, to provide the correct matrix scoring. I’m happy to announce that the twenty or so corrections made yesterday were added to the 120,000 or so corrections made over the past ten years. With these corrections the LRT gets better and stronger every week. Minimizing taxon exclusion maximizes the opportunity to correctly nest new and enigma taxa with old and established taxa, even if the new and old specimens are incomplete or scattered about.

The earlier August 2017 blogpost for Fruitafossor
was updated yesterday to erase old errors and enter the corrections.


References
Huttenlocker AD, Grossnickle DM, Kirkland JI, Schultz JA and Luo Z-X 2018. Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana. Nature Letters  Link to Nature
Luo Z-X and Wible JR 2005. A late Jurassic digging mammal and early mammal diversification. Science 308:103–107.
Shaw G 1792. Musei Leveriani explicatio, anglica et latina.

wiki/Fruitafossor
digimorph.org/specimens/Fruitafossor_windscheffeli/

 

New genomic estimate misses monotreme-marsupial split by 43 million years

Summary for those in a hurry:
Fossils provide hard evidence. Deep time gene studies provide estimates and false positives too often to trust them.

Zhou et al. 2021 report:
“Our phylogenomic reconstruction shows that monotremes diverged from therians around 187 million years ago, and the two monotremes diverged around 55 million years ago. This estimate provides a date for the monotreme–therian split that is earlier than previous estimates (about 21 million years ago, but agrees with recent analyses of few genes and fossil evidence.”

Let’s stop putting our faith in estimates derived from genomic deep time studies that have proven themselves to be wrong too many times. Here, the Zhou et al. estimate is at least 43 million years too late (Fig. 2) based on Brasilitherium (Fig. 3) fossils and the tree topology recovered by the LRT (Fig. 1).

Figure 7. Subset of the LRT focusing on Metatheria (marsupials) including Paedotherium and Adalatherium.

Figure 1. Subset of the LRT focusing on Metatheria (marsupials) including Paedotherium and Adalatherium.

By contrast with Zhou et al. 
Morganucodon (Late Triassic, 205mya, Fig. 4) is a basal marsupial in the large reptile tree (LRT, 1790+ taxa; subset Fig. 1) based on phenomic (= trait) analysis that includes fossil taxa. Genomic tests are infamous for false positives when dealing with deep time taxa.

Figure 1. Select basal cynodonts and mammals set chronologically. The divergence times for placentals (Eutheria), marsupials (Metatheria) and monotremes (Mammalia) are estimated here.

Figure 2. Select basal cynodonts and mammals set chronologically. The divergence times for placentals (Eutheria), marsupials (Metatheria) and monotremes (Mammalia) are estimated here.

Brasilitherium,
(Figs. 3, 4) from the Early Norian, Late Triassic, 225mya, is a derived monotreme in the LRT. That means it lived AFTER the monotreme-therian split which must have occurred at least 230mya.

Figure 1. Brasilodon compared to Kuehneotherium, Akidolestes and Ornithorhynchus, the living platypus.

Figure 3. Brasilodon compared to Kuehneotherium, Akidolestes and Ornithorhynchus, the living platypus.

As everyone knows
the platypus and echidna are highly derived monotremes. Megazostrodon (Fig. 4) is the last common ancestor (LCA) of all monotremes and all mammals. Megazostrodon was a Late Jurassic late survivor of that earlier (Middle Triassic?) radiation.

Figure 5. Basal mammals and their proximal ancestors. Here taxa below Megazostrodon are mammals. Those above are not. Hadrocodium is uniquely reduced, but this occurs within the Mammalia.  The dual jaw joint was tentatively present in Pachygenelus.

Figure 4. Basal mammals and their proximal ancestors. Here taxa below Megazostrodon are mammals. Those above are not. Hadrocodium is uniquely reduced, but this occurs within the Mammalia.  The dual jaw joint was tentatively present in Pachygenelus.

According to the LRT,
there was no gradual ascent of monotremes leading to marsupials. Rather the monotreme-marsupial split occurred at the origin of mammals and monotremes. How this affects the genes for lactation discussed in the Zhou et al. paper is beyond the scope of this blogpost.

The purpose here
is to emphasize the importance of a broad, proper and valid phylogenetic context before proceeding to the narrow focus of your interests. 42 co-authors using cutting edge genomic techniques hobbled their otherwise excellent and in-depth report by skipping step number one.


References
Zhou Z et al. (41 co-authors) 2021. Platypus and echidna genomes reveal mammalian biology and evolution. Nature https://doi.org/10.1038/s41586-020-03039-0

 

Marsupial cladograms: Tooth traits recover false positives

A traditional dependence on molar traits
has obscured and mixed up fossil mammal relationships (Fig. 1) when compared to phenomic studies using skulls and skeletons of extant taxa and fossils (Fig. 2). In this way, tooth traits are shown to be like gene traits. They deliver false positives that are not validated by taxa tested by a large suite of traits from nose to tail.

Luo et al. 2011
published their cladogram nesting Juramia as the basalmost placental recovered from a cladogram of mammal interrelations based largely on tooth traits (Fig. 1).

Figure 1. From Luo et al. 2011, mammal claodgram focused on Juramaia employing many tooth traits. Compare to the LRT in figure 2 which minimizes tooth traits.

Figure 1. From Luo et al. 2011, mammal claodgram focused on Juramaia employing many tooth traits. Compare to the LRT in figure 2 which minimizes tooth traits.

In stark contrast,
the large reptile tree (LRT, 1728+ taxa) nested Juramaia as a monotreme (Prototheria).

Figure 4. Subset of the LRT cladogram of basal Mammalia. Note the traditional clade Metatheria is a grade with new names proposed here.

Figure 2. Subset of the LRT cladogram of basal Mammalia. Note the traditional clade Metatheria is a grade with new names proposed here.

Figure 2. Juramaia (Late Jurassic, 160 mya) is more completely known and nests between monotremes and therians (marsupials + placentals).

Figure 3. Juramaia (Late Jurassic, 160 mya) is more completely known and nests with monotremes not placentals.

 

References
Luo Z-X, Yuan C-X, Men Q-J and JiQ 2011. A Jurassic eutherian mammal and divergence of marsupials and placentals. Nature 476: 442–445. doi:10.1038/nature10291.

Asiatherium enters the LRT: mammal nomenclature issues follow

Everyone agrees
that Asiatherium (Figs, 1,2) nests close to Monodelphis, Caluromys and placentals. Trofimov and Szalay 1994 agreed. So did Denyer, Regnault and Hutchinson 2020. So did the large reptile tree (LRT, 1729+ taxa, subset Fig. 3).

Figure 1. Asiatherium in situ from Szalay and Trofimov 1996.

Figure 1. Asiatherium in situ from Szalay and Trofimov 1996.

Asiatherium reshetovi (Trofimov and Szalay 1994, Szalay and Trofimov 1996; PIN 3907; Late Cretaceous; 80mya; Figs. 1, 2) is a key Mongolian metathere ancestral to monodelphids and Caluromys, which is ancestral to placentals. It is derived from Triassic sisters to extant late survivors, DidelphisGilronia and Marmosops.

Figure 2. Asiatherium skull slightly modified from Szalay and Trofimov 1996. Colors added here.

Figure 2. Asiatherium skull slightly modified (longer lateral view premaxilla to match dorsal and ventral views) from Szalay and Trofimov 1996. Colors added here.

The problem is,
according to results recovered by the LRT, mammal clade nomenclature needs to go back to basics. Several modern mammalian clade names are found to be junior synonyms of traditional clades in the LRT.

Prototheria (Gill 1872) is a junior synonym
for Monotremata (Bonaparte 1837) in the LRT.

According to Wikipedia, “Prototheria is a paraphyletic subclass to which the orders MonotremataMorganucodontaDocodontaTriconodonta and Multituberculata have been assigned, although the validity of the subclass has been questioned.”

In the LRT Morganucodon is a a marsupial (see below). Docodon is a taxon within Monotremata. Triconodon is a taxon within Monotremata. Multituberculata is a clade within the placental clade Glires (Fig. 4). So, the clade Monotremata is monophyletic and has precedence.

Theria (Parker and Haswell 1897) is a junior synonym
of Marsupialia (Illiger 1811). Metatatheria (Thomas Henry Huxley 1880) is also a junior synonym of Marsupialia.

The late-surviving basalmost marsupial in the LRT (Fig. 4), Ukhaatherium (Fig. 3), has epipubic (marsupial) bones. That long rostrum indicates this taxon is close to monotremes.

Figure 3. Ukhaatherium in situ.

Figure 3. Ukhaatherium in situ.

Unlike the monophyletic clade Monotremata,
a series of nested marsupial clades are present. The last of these gives rise to Placentalia, only one of several that lose the pouch (Fig. 4). New names are proposed here where appropriate:

  1. Marsupialia = Ukhaatherium and kin + all descendants (including placentals)
  2. Paleometatheria = Morganucodon and kin + all descendants.
  3. Didelphimetatheria = Eomaia and kin + all descendants
  4. Phytometatheria = Marmosops and kin + all descendants
  5. Carnimetatheria = Asiatherium and kin + all descendants
  6. Transmetatheria = Caluromys and kin + all descendants
  7. Placentalia = Vulpavus and kin + all descendants
Figure 4. Subset of the LRT cladogram of basal Mammalia. Note the traditional clade Metatheria is a grade with new names proposed here.

Figure 4. Subset of the LRT cladogram of basal Mammalia. Note the new names proposed here.

Basal marsupial taxa are omnivores. 
Derived phytometatheres are herbivores. Derived carnimetatheres are carnivores to hyper-carnivores. Transmetatheres (Carluromys) and basal Placentalia remain omnivores.

In the LRT Eutheria (Gill 1872) is a junior synonym
of Placentalia (Owen 1837). Omnivorous civets like Nandinia are basal placentals. Carnivora is a basal placental clade following basal placental civets.

Competing cladograms
Denyer, Regnault and Hutchinson 2020 recently looked at the marsupial patella, or more specifically the widespread absence or reduction of the kneecap. The authors concluded, “metatherians independently ossified their patellae at least three times in their evolution.”

Unfortunately, Denyer et al. tested Caenolestes, the ‘shrew opossum’. Not surprisingly it nested close to placentals in their cladogram. Caenolestes was earlier nested in the LRT within the placental clade, Glires, closer to shrews than to opossums. It has no pouch, but converges with marsupials in several aspects. Inappropriate taxon inclusion, like Caenolestes, occurs due to taxon exclusion. Excluded taxa would have attracted and removed the inappropriate taxon. Taxon exclusion plagues Denyer et al.

Historically, you may remember,
Bi et al. 2018, while presenting Early Cretaceous Ambolestes, suffered from massive taxon exclusion and traditional bias in attempting to produce a cladogram of mammals. Bi et al. recovered Sinodelphys (Early Cretaceous) and Juramaia (Late Jurassic) as ‘eutherians’. In the LRT both are monotremes.

Other basal mammal cladograms
depend too much on tooth traits. Convergence in tooth traits creates problems, as documented earlier. We’ll look at this problem in more detail soon.

The above subset of the LRT appears to be a novel hypothesis
of interrelationships. If not, please provide a citation so I can promote it.


References
Bi S, Zheng X, Wang X, Cignetti NE, Yang S, Wible JR. 2018. An Early Cretaceous eutherian and the placental marsupial dichotomy. Nature 558(7710):390395 DOI 10.1038/s41586-018-0210-3.
Denyer AL, Regnault S and Hutchinson JR 2020. Evolution of the patella and patelloid in marsupial mammals. PeerJ 8:e9760 http://doi.org/10.7717/peerj.9760
Szalay FS and Trofimov BA 1996. The Mongolian Late Cretaceous Asiatherium, and the early phylogeny and paleogeography of Metatheria. Journal of Vertebrate Paleontology 16(3):474–509.
Trofimov BA and Szalay FS 1994. New Cretaceous marsupial from Mongolia and the early radiation of Metatheria. Proceedings of the National Academy of Sciences 91:12569-12573

Cladogram of the Mammalia (subset of the LRT)

A summary today…
featuring a long cladogram (Fig. 1), a subset from the large reptile tree (LRT, 1259 taxa) focusing on the Mammalia. This is how this LRT subset stands at present. Not much has changed other than the few node changes from the past week.

The transition from Prototheria to Theria (Metatheria)
includes long-snouted taxa, like Ukhaatherium. Nearly all Prototheria are also long-snouted (Cifelliodon is the current sole exception).

The transition from Metatheria to Eutheria (simplified)
includes small omnivorous didelphids arising from the carnivorous/herbivorous split among larger metatherians. Basal Carnivora, the most basal eutherian clade, are also omnivores. Caluromys, the extant wooly opossum, has a pouch, but nests at the base of all placental taxa (the LRT tests only skeletal traits), so it represents the size and shape of the earliest placentals (contra O’Leary et al. 2013)… basically didelphids without pouches, and fewer teeth, generally (but not always).

Basal members of most placental clades
are all Caluromys-like taxa, with a rapid radiation in the Late Triassic/Early Jurassic generating most of the major placental clades in the LRT (Fig. 1). Larger members of each of these placental clades appeared in the fossil record only after the K-T extinction event. So hardy where these basal taxa, that many still live to this day.

As shown earlier, higher eutheria are born able to able to walk or swim. They are no longer helpless with arboreal parents (tree-climbing goats the exception). Basal eutherians reproduce more like their metatherian ancestors, with helpless infants.

Figure 1. Subset of the LRT focusing on mammals.

Figure 1. Subset of the LRT focusing on mammals. Extant taxa are colored. Thylacinus is recently extinct.

The latest competing study
(O’Leary et al. 2013, Fig. 2) recovers the highly specialized edentates, aardvarks, elephants and elephant shrews as the most primitive placentals. Carnivora + bats are quite derived in the O’Leary team cladogram, somehow giving rise to ungulates and whales. This is an untenable hypothesis. It doesn’t make sense. Evidently the O’Leary team had faith that smaller didelphid-like ancestors would fill in the enormous phylogenetic gaps in their cladogram. By contrast the LRT has all the operational taxonomic units (OTUs) it needs to produce a series of gradually accumulating derived traits between every taxon in its chart (Fig. 1). The LRT makes sense.

Figure 5. Simplified version of the O'Leary et al 2013 cladogram showing placental relations exploded after the K-T boundary.

Figure 5. Simplified version of the O’Leary et al 2013 cladogram showing placental relations exploded after the K-T boundary.

References
O’Leary, MA et al. 2013. The placental mammal ancestor and the post-K-Pg radiation of  placentals. Science 339:662-667. abstract
Wible JR, Rougier GW, Novacek MJ, Asher RJ 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary Nature 447: 1003-1006

https://pterosaurheresies.wordpress.com/2016/08/31/another-look-at-the-oleary-et-al-hypothetical-ancestor-of-placentals/

https://pterosaurheresies.wordpress.com/2013/02/15/post-k-t-explosion-of-placentals-oleary-et-al-2013/

ArchibaldEtAl.pdf
protungulatum-donnae website

Chaoyangodens: a transitional monotreme with big canines

Hou and Meng 2014
described a new Jehol eutriconodont mammal, Chaoyangodens lii, (Fig. 1) from the Yixian formation, Early Cretaceous. “The new species has a tooth formula I5-C1-P1-M3/i4-c1-p1-m4, unique among eutriconodonts in having only one premolar in lower and upper jaws, respectively, and a distinctive diastema between the canine and the premolar. Its simple incisors and reduced premolars show a mosaic combination of primitive and derived features.” In other words, this is a transitional taxon, as most are.

Later, Meng and Hou 2016
described ‘the earliest known mammalian stapes’ from the same specimen. “The stapes of Chaoyangodens is reduced in size compared to those of non-mammalian cynodonts and is within the size range of extant mammals.”

Figure 1. Chaoyangodens lii in situ and restored skull in lateral view. This taxon is a monotreme basal to both the echidna and platypus.

Figure 1. Chaoyangodens lii in situ and restored skull in lateral view. At a screen resolution of 72 dpi, this image is twice life size. This mouse-sized taxon is a monotreme basal to both the echidna and platypus.

Figure 2. Subset of the LRT focusing on monotremes and Chaoyangodens.

Figure 2. Subset of the LRT focusing on monotremes and Chaoyangodens.

Here in the large reptile tree (LRT, 1137 taxa, Fig. 2) Chaoyangodens nests between Kuehneotherium and Akidolestes, basal  to the living monotremes, Ornithorhynchus and Tachyglossus.

The top of the Chaoyangodens skull is buried in the matrix. The shape of the skull in lateral view, or at least parts of it, like the position of the orbit (Fig. 1), can be surmised by phylogenetic bracketing.

Based on the nesting of Chaoyangodens
and relatives, like Brasilitherium and Kuehneotherium (Late Triassic), these taxa are all crown mammals, not stem mammals (contra traditional thinking).

Luo, Kielan-Jaworowska and Cifelli (2002)
also nested eutriconodonts within crown mammals and this was confirmed by many later workers. The LRT nests many traditional triconodonts and eutriconodonts elsewhere, both more primitive and more derived.

Recently
we looked at the echidna sister/ancestor, Cifelliodon, here. It also had fewer and bigger teeth in the jaws, though none of those erupted beyond the gum line.

References
Hou S-L and Meng J 2014. A new eutriconodont mammal from the Early Cretaceous Jehol Biota of Liaoning, China. Chinese Science Bulletin 59, 546–553.
Luo Z-X, Kielan-Jaworowska  z and Cifelli RL 2002. In quest for a phylogeny of Mesozoic mammals. Acta Palaeontologica Polonica. 47 (1): 1–78.
Meng J and Hou S-L 2016. Earliest known mammalian stapes from an early Cretaceous eutriconodontan mammal and implications for evolution of mammalian middle ear. Palaeontologica Polonica 67:181–196.

wiki/Eutriconodonta

Cifelliodon: new echidna ancestor from the Early Cretaceous of Utah

This one seemed pretty obvious from the first impression,
but failed to make the same impression on the original authors (Huttenlocker et al., 2018).

Usually mammal teeth are found without a skull.
Huttenlocker et al., 2018 found a skull largely without teeth (most don’t erupt beyond the rim of the very few alveoli), certainly a derived trait for mammals. And this is one more way tetrapods became toothless. They named their new taxon, Cifelliodon wahkarmoosuch (Fig. 1).

Figure 1. Early Cretaceous Cifelliodon is ancestral to the living echidna, Tachyglossus according to the LRT. The lack of teeth here led to toothlessness in living echidnas. The skull of Tachyglossus is largely fused together, lacks teeth and retains only a tiny lateral temporal fenestra (because the jaws don't move much in this anteater. Compared to Cifelliodon the braincase is greatly expanded, the lateral arches are expanded and the two elements fuse, unlike most mammals.

Figure 1. Early Cretaceous Cifelliodon is ancestral to the living echidna, Tachyglossus according to the LRT. The reduced number and size of teeth here led to toothlessness in the living echidna. The skull of Tachyglossus retains only a tiny lateral temporal fenestra (because the jaws don’t move much in this anteater. Compared to Cifelliodon the braincase is greatly expanded, the lateral arches are expanded and the two elements fuse, unlike most mammals.

According to Wikipedia:
“Cifelliodon is an extinct genus of haramiyid mammal from the Lower Cretaceous of North America. It is a mammaliaform, and is one of the latest surviving haramiyids yet known, belonging to the family Hahnodontidae. Its discovery led to the proposal to remove hahnodontids from the larger well-known group, the multituberculates.”

As usual the LRT recovered a different nesting.

Figure 2. Cifelliodon skull in three views, plus DGS, plus the original drawing, which is not very accurate.

Figure 2. Cifelliodon skull in three views, plus DGS, plus the original drawing, which is not very accurate. A mandible is restored here.

Figure 3. Subset of the LRT focusing on Monotremes, now including Cifelliodon.

Figure 3. Subset of the LRT focusing on Monotremes, now including Cifelliodon.

Here
in the large reptile tree (LRT, 1233 taxa) Cifelliodon wahkarmoosuch from the Early Cretaceous of Utah nests strongly with Tachyglossus (Figs. 1, 4, 5), one of the extant egg-laying echidnas, currently restricted to Australia and surrounding islands. Tachyglossus was tested in the (Huttenlocker et al. analysis of basal mammal relationships, but the two taxa did not nest together.

Unfortunately Huttenlocker et al., 2018
experienced taxon exclusion problems that nested Cifelliodon with the distinctly different wombat Vintana and those two with the distinctly different multituberculates all more primitive than monotremes.

To their credit
Huttenlocker et al. linked this North American taxon with others from Gondwana which includes Australia, which broke off 99 mya, 40 million years after the appearance of Cifelliodon in Utah. In an interview for USC, Huttenlocker reported, “Most of the fossil record of early mammal relatives is based on teeth. Cifelliodon is unique in that it is one of the only near-complete skulls of a mammal relative from the basal Cretaceous of North America and is the only fossil of early mammal relatives from this time interval in Utah.”

“The fact that the skull looked so primitive compared to other known mammal groups from the Cretaceous made figuring out its relationships extremely difficult. It shows some unique dietary specializations that are seen in only a handful of groups that lived during the age of dinosaurs. Ultimately, the structure of the preserved molars showed clear similarities to some neglected fossil teeth from Northern Africa. So we think that Cifelliodon represents an archaic offshoot whose relatives may have dispersed into the southern continents and became fairly successful during the Cretaceous.”

Figure 3. Tachyglossus skeleton, manus and x-rays. Note the perforated pelvis.

Figure 4. Tachyglossus skeleton, manus and x-rays.

The skull of Tachyglossus
retains only a tiny lateral temporal fenestra (because the jaws don’t move much in this anteater. Compared to Cifelliodon the braincase is greatly expanded, the lateral arches are expanded and the two elements fuse, unlike most mammals.

Figure 1. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws.

Figure 5. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws. Many of the derived traits seen here developed during the last 100 million years since Cifelliodon.

 

References
Huttenlocker AD, Grossnickle DM, Kirkland JI, Schultz JA and Luo Z-X 2018. Late-surviving stem mammal links the lowermost Cretaceous of North America and Gondwana. Nature Letters  Link to Nature

wiki/Cifelliodon

https://news.usc.edu/143411/why-you-should-care-about-this-130-million-year-old-fossil/

Tachyglossus, the other egg-laying mammal

Figure 1. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws.

Figure 1. The echidna (genus: Tachyglossus) in life. This slow-moving spine-covered anteater has digging claws.

Tachyglossus aculeatus (Shaw 1792) is the echidna and the only other genus of egg-laying mammal. It protects itself with sharp spines and has a long, ant-catching tongue. The hands and feet are adapted to digging with short, almost immobile proximal elements (Fig. 3) and long claws. Prepubic bones precede the pubis. A proximal process sits atop the fibula. The leathery snout without whiskers is sensitive to vibrations.

Figure 2. The skull of Tachyglossus is largely fused together, lacks teeth and has no lateral temporal fenestra (because the jaws don't move much in this anteater.

Figure 2. The skull of Tachyglossus is largely fused together, lacks teeth and has no lateral temporal fenestra (because the jaws don’t move much in this anteater. Hard to find sutures here. Let me know if you have better data to make corrections.

Distinct for its sister,
Ornithorhynchus, and many other mammals, the acetabulum is perforated. The lateral temporal fenestra is absent. So are the teeth. Like the hedgehog, the echidna can roll itself into a ball for protection.

Figure 3. Tachyglossus skeleton, manus and x-rays. Note the perforated pelvis.

Figure 3. Tachyglossus skeleton, manus and x-rays. Note the perforated pelvis.

There are those
who say characters define a taxon. We have to get away from that hypothesis. Here a perforated acetabulum would make Tachyglossus a dinosaur, to the late Larry Martin’s delight. Tachyglossus has no temporal fenestra. So, does that make it an anapsid? No. The only thing that tells us what a taxon is… is its placement on a wide gamut cladogram that tests hundreds of candidate sister taxa and hundreds of traits. Testing a suite of several hundred traits in a wide gamut study is the only way to confidently determine taxonomy and avoid the pitfalls of convergence and taxon exclusion that plague smaller studies that too often fail to minimize false positives and ‘by default’ nestings. And some DNA studies cannot be validated, except by morphological studies.

References
Shaw G 1792. Musei Leveriani explicatio, anglica et latina.

wiki/Tachyglossus

There’s nothing special about Henosferus

The incisors are not too big
or weird or crowded (Fig. 1), the canine just rises above the rest of the teeth, there are only 5 premolars all standard-shaped, and only three molars, all standard-shaped. The dentary definitely formed the main jaw joint and the post-dentary bones must have been tiny.

Figure 1. Henosferus mandible restored by Rougier et al. 2005 from several broken specimens.

Figure 1. Henosferus mandible restored by Rougier et al. 2005 from several broken specimens.

…and that’s why
Henosferus ( Rougier et al. 2007; Middle Jurassic) makes a good candidate for basalmost mammal. There are too few traits here to add it to the large reptile tree (LRT). Frankly, I’m eyeballing this restoration. It compares well with Juramaia (Fig. 2) without the odd molars and incisors. 

Figure 2. Juramaia (Late Jurassic, 160 mya) is more completely known and nests between monotremes and therians (marsupials + placentals).

Figure 2. Juramaia (Late Jurassic, 160 mya) is more completely known and nests between monotremes and therians (marsupials + placentals).

Henosferus is traditionally considered
a member of the Australosphenida, a group of mammals that include monotremes, and other taxa known chiefly from scraps. Vincelestes sometimes makes this list, but in the LRT it nests as a carnivorous marsupial.

References
Luo Z-X, Yuan C-X, Men Q-J and JiQ 2011. A Jurassic eutherian mammal and divergence of marsupials and placentals. Nature 476: 442–445. doi:10.1038/nature10291.
Rougier, GW, Martinelli AG, Forasiepi AM and Novacek M J 2007. New Jurassic mammals from Patagonia, Argentina : a reappraisal of australosphenidan morphology and interrelationships. American Museum novitates, no. 3566. online here.

wiki/Juramaia
wiki/Henosferus

Docodon and molar count

The genus Docodon (Marsh 1881; Late Jurassic, 170 mya; Fig. 1) is represented by a jaw with several more molars than typical (Fig. 1). Hard to tell the premolars from the molars in lateral view. See the dorsal view for the distinct difference. Even so, the count may be off, because molars are not molars based on shape, but on the fact that they appear once and are not replaced during growth. I cannot tell, note have I found references that say where the division is in Docodon.

Figure 1. The holotype of Docodon has 4 incisors, 4 premolars and a whopping 7 molars.

Figure 1. The holotype of Docodon has 4 incisors, 4 premolars and a whopping 7 molars. diplocynodon has 8 according to Osborn, who confirms the 7 in Docodon.

More than four molars in a mammal jaw is relatively rare.
But not rare in ancestral monotremes. Among tested taxa  Amphitherium, Kuehneotherium and Akidolestes have 6.

Figure 1. The addition of teeth in Kuehneosaurus and Akidolestes led to the loss of teeth in Ornithorhynchus.

Figure 1. The addition of teeth in Kuehneosaurus and Akidolestes led to the loss of teeth in Ornithorhynchus.

There is also “a rule” that says
only one canine appears, but the other teeth can vary greatly in number. I’m wondering if that is true. Sometimes there will be a small, simple tooth arising between the canine and the double-rooted premolars. Is that a tiny canine? or a tiny premolar? Maybe someone out there has not only the answer, but the reason why.

And yes,
I’m aware of the convention that numbers premolars 1-4. But the anterior one, is almost always the smallest, as if it just arrived.

References
Marsh OC 1881. Notice of new Jurassic mammals. American Journal of Science. (3) xxi: 511-513.

wiki/Docodon