Ever hear of Palaeothentes?

Currently there is no Wikipedia page for this taxon.
Even so, I found it to be far more important at filling gaps and shaking up paradigms than it seemed at first. The small, dull-looking taxa tend to be like that, as readers now know.

You could find this rat-sized Miocene taxon
(Fig. 1) in Carroll’s 1988 book, Vertebrate Paleontology, now well-worn and in pieces due to constant page flipping and scanning. Today’s research has revealed several more precise and more recent resources.

Figure 1. Not a marsupial, and not a shrew opossum, Palaeothentes nests in the LRT at the base of the Apatemys + Trogosus clade nest to the clade of living shrew opossums within Glires.

Figure 1. Not a marsupial, and not a shrew opossum, Palaeothentes nests in the LRT at the base of the Apatemys + Trogosus clade next to the clade of living shrew opossums within Glires.

According to Abello and Candella 2010,
Palaeothentes minutes (Ameghino 1887) is a paucituberculatan (details below) from the Santa Cruz Formation The results indicate that Palaeothentes would have been an agile cursorial dweller, with leaping ability, similar to the extant paucituberculatan Caenolestes fuliginosus and the didelphid Metachirus nudicaudatus.”

Okay, so now we have a problem.
In the large reptile tree (LRT, 1445 taxa) Caenolestes is not a marsupial. It nests with Rhyncholestes and more distantly Apatemys and more distantly, the extant tree shrew, Tupaia and the extant shrew, Scutisorex. As noted earlier, shrew opossums are placental shrews, not marsupial opossums in the LRT.

Wikipedia reports,
“Like several other marsupials, they do not have a pouch, and it appears that females do not carry the young constantly, possibly leaving them in the burrow.”
That’s describes most rodent/rabbit/tree shrew mothers and their young.

Wikipedia reports,
“Paucituberculata is an order of South American marsupials. Although currently represented only by the eight living species of shrew opossums, this order was formerly much more diverse, with more than 60 extinct species named from the fossil record, particularly from the late Oligocene to early Miocene epochs.”

Let’s solve that problem
by adding Palaeothentes to the LRT. Doing so recovers this taxon at the base of the Apatemys + Trogosus clade, next to the clade that includes Caenolestes, within Glires, far from Marupialia.

I suspect taxon exclusion
is the cause for the present lack of confirmation for traditional consensus. Many PhDs over several decades have followed tradition in nesting and testing shrew opossums with marsupials without testing them against apatemyids apparently. That’s why the LRT is here, to test taxa that have never been tested together before.

But wait! There’s a novel twist here~~~~~~!
Carroll 1988 reports, “Caenolestids have long been recognized as being very distinct from other South American marsupials, but they share with them a highly distinctive pattern of the spermatozoa, which become paired within the epididymis. Paired sperm are not known in any placental groups or among the Australian marsupials.” 

Sorry.
Physical traits have to trump genes and sperm. It just has to be that way because the LRT includes fossil taxa, which never preserve sperm. There have to be rules that all participants abide by. Interesting that the gene for paired spermatozoa is localized to one continent, just as genes separate other placentals into afrotheres and laurasiatheres. By the way, “The data show that paired spermatozoa exhibit a significant motility advantage over single spermatozoa in a viscous medium” according to Moore and Taggart 1995, who tested Monodelphis, a South American opossum.

Finally
we have a last common ancestor for arboreal Apatemys (Eocene, North America) and terrestrial Trogosus (Eocene, North America), two former enigma taxa with little to no relationship with other better known mammal clades. All members of Glires had their genesis sometime in the Jurassic, based on the presence of highly derived multituberculates (clade: Glires) in the Jurassic.

Wikipedia considers
apatemyids and trogosinae (Tillodontia) to be members of the Cimolesta, “an extinct order of non-placental eutherian mammals.” This bungling of the mammal family tree is due to taxon exclusion and the lack of a phenomic (trait-based) wide gamut cladogram that includes all the taxa present in the LRT. Paleontology needs to toss off a wide range of useless tradition with a reptile revolution led by someone out there confirming (or refuting) the widest gamut cladogram presently available: the LRT.


Palaeothentes lemoinei (Ameghino 1887, MPM-PV 3566; Miocene) was considered a prehistoric shrew opossoum (clade: Paucituberculata) but here nests next to shrew opossums, at the base of the Apatemys + Trogosus clade within Glires. The skull is 2x wider than tall, the canines are still large, the last premolar is large with a flat occlusal surface and the nasals split to form a zigzag suture with the frontals.


References
Abello MA and Candela AM 2010. Postcranial skeleton of the Miocene Marsupial Palaeothentes(Paucituberculata, Palaeothentidae): Paleobiology and Phylogeny. Journal of Vertebrate Paleontology 30(5):1515-1527.
Ameghino F 1887. Enumeracions sistematicad e las especies de mamiferos
fosiles coleccionados por Carlos Ameghino en los terranos eocenos de la Patagonia austral y depositados en el Museo La Plata. Boletin Museo de La Plata, 1:1-26.
Carroll RL 1988. Vertebrate Paleontology and Evolution. W. H. Freeman and Co. New York.
Forasiepi AMSánchez-Villagra MR, Schmelzle T,  Ladevèze S and Kay RF 2014. An exceptionally well-preserved skeleton of Palaeothentes from the Early Miocene of Patagonia, Argentina: new insights into the anatomy of extinct paucituberculatan marsupials. Swiss Journal of Palaeontology, 133(1):1-21.
Moore HD and Taggart DA 1995. Sperm pairing in the opossum increases the efficiency of sperm movement in a viscous environment. Biol. Reprod. 52(4):947-53.
Osgood WH 1921. A monographic study of the American marsupial, Caenolestes. Field Museum of Natural History, Zoological series 14:1–156.

wiki/Apatemyidae
wiki/Paucituberculata
https://en.wikipedia.org/wiki/Shrew_opossum
wiki/Vertebrate_Paleontology_and_Evolution

Paucituberculata -Trouessart 1898, Ameghino 1894


There was some news
about Palaeothentes recently (see below). Note, the experts consulted here consider this genus a marsupial.

New Bolivian Marsupials from the Middle Miocene

Two new Royal Society papers suffer from taxon exclusion

Gutarra et al. 2019
tested the effects of several body plans on the hydrodynamic drag of simplified 3D digital ichthyosaurs. They reported, “Our results show that morphology did not have a major effect on the drag coefficient or the energy cost of steady swimming through geological time.”

Unfortunately
the Gutarra team included the basal sauropterygian ichthyosaur-mimic Cartorhynchus as their basal taxon, ignoring the following four valid ichthyosaur basal taxa.

  1. Wumengosaurus
  2. any hupehsuchid
  3. Thaisaurus
  4. Xinminosaurus

Given the Gutarra et al. similar results
for all included digitally generated taxa, it would have been instructive to test at least one of these basal taxa or perhaps outgroup taxa from the Mesosauria and/or Thalattosauria in order to set a baseline. Co-author professor MJ Benton has been reprimanded for excluding taxa several times before, and doggone it, he did it again.


Halliday et al. 2019
“supports a Late Cretaceous origin of crown placentals with an ordinal-level adaptive radiation in the early Paleocene, with the high relative rate permitting rapid anatomical change without requiring unreasonably fast molecular evolutionary rates.” 

By contrast
the large reptile tree (LRT, 1413 taxa) nests several placental taxa (like multituberculates) in the Jurassic with placental origins likely in the Late Triassic very soon after the origin of Mammalia.

Halliday’s team differentiates extant placentals from several extinct eutherians,
while the LRT finds only one extant taxa, the arboreal didelphid Caluromys, in the Eutheria outside of the Placentalia.

Halliday’s team cites the Luo et al. 2011 report
of “a Jurassic eutherian mammal” (= Juramaia) with reservations. In the LRT Juramaia nests with basal prototherians, not eutherians.

None of Halliday’s published work
matches the topology of the LRT. The Halliday team nests highly derived hedgehogs, elephants and armadillos as a closely related clade at the base of their cladogram of extant placentals.

By contrast and employing more taxa
the LRT documents the evolution of three clades of basal placentals, like arboreal civets, bats, dermopterans, pangolins and tree shrews (Primates + Glires), from arboreal marsupials, like Caluromys. 

Evolution: small changes over time.
The editors and referees approved Halliday’s ‘traditional’ topology. Someone should have checked results for relationships that minimize differences between recovered sisters. More taxa and avoiding genetic scoring would have helped.

Halliday’s study supports several invalidated genetic clades,
including Atlantogenata (anteaters + elephants and kin), Boreotheria (mice + whales + humans and kin), and Afrotheria (elephant shrews + elephants and kin). Even so, editors, paleoworkers and referees approved these untenable and refuted relationships.

That’s why the LRT is here,
to lift the covers and show you untenable traditional relationships, then to offer a tree topology in which all included taxa document a gradual accumulation of derived traits.


References
Gutarra S, Moon BC, Rahman IA, Palmer C, Lautenschlager S, Brimacombe AJ, and Benton MJ 2019. Effects of body plan evolution on the hydrodynamic drag and energy requirements of swimming in ichthyosaurs. Proc. R. Soc. B 286: 20182786. http://dx.doi.org/10.1098/rspb.2018.2786
Halliday TJD, dos Reis M, Tamuri AU, Ferguson-Gow H, Yang Z and Goswami A 2019. Rapid morphological evolution in placental mammals post-dates the origin of the crown group. Proc. R. Soc. B 286: 20182418. http://dx.doi.org/10.1098/rspb.2018.2418

Hey! Some of those Miocene ‘ungulates’ are marsupials!

More heresy today
courtesy of taxon inclusion.

Cassini 2013
looked at several traditional Miocene South American ‘ungulates’ (Fig. 1) unaware that these taxa do not nest in monophyletic clade any more specific than Theria in the large reptile tree (LRT, 1401 taxa). Cassini was reporting the results of “an ecomorphological study based on geometric morphometrics of the masticatory apparatus.” So he was working from prior cladograms and focusing on the mechanics of eating.

Figure 1. Image from Cassini 2013. Pink taxa are marsupials. Others are placentals.

Figure 1. Image from Cassini 2013. Pink taxa are marsupials. Others are placentals.

Earlier
here and here the traditional clade Notoungulata was splintered by the LRT into several clades, some among the marsupials, other among the placentals.

Traditional ‘Litopterna’
Diadiaphorus (Fig. 1) nests at the base of this clade. Theosodon (Fig. 1) nests as a derived taxon. Also included, but not listed: Chalicotherium and other chalicotheres.

Considering its member taxa,
the clade Litopterna (Ameghino 1889) is a junior synonym for Chalicotheridae (Gill 1872) in the LRT.

Considering its member taxa,
the clade Astrapotheria (Lydekker 1894) is a junior synonym for Meniscotheriinae (Cope 1882) and both nest within Phenacodontidae (Cope 1881).

Interatheriidae (Ameghino 1887) traditionally includes Interatherium, Protypotherium, Miocochilius and other taxa listed here. In the LRT Interatherium nests close to the ancestry of the Toxodon clade + the kangaroo clade within Metatheria. By contrast, Protypotherium and Miocochillus nest with Homalodotherium deep within the Eutheria. Homalodotherium traditionally nests with the the metatherian Toxodon. According to the LRT, all the above taxa developed similar enough traits by convergence that all were mistakenly lumped together in the invalid placental clade ‘Notoungulata’.

This is not the first time
metatherians were split from convergent eutherians. Most creodonts are marsupial predators, phylogenetically distinct from their traditional sisters in the clade Carnivora, within the clade Eutheria (Placentalia).

New taxa added to the LRT:

  1. Hegetotherium (Fig. 1) nests with Mesotherium between Interatherium and the Toxodon clade in the Metatheria. 
  2. Diadiaphorus (Fig. 1), the horse-mimic, nests at the base of the Litopterna/Chalicotheriidae, just basal to Litolophus.

I did not know these two, so I added these two to better understand them.

References
Cassini G 2013. Skull Geometric Morphometrics and Paleoecology of Santacrucian (Late Early Miocene; Patagonia) Native Ungulates (Astrapotheria, Litopterna, and Notoungulata). Ameghiniana 50 (2):193–216. DOI: 10.5710/AMGH.7.04.2013.606

Hapalodectes: when primates split from dolphins

Back when placental mammals were first diversifying in the Jurassic
they all looked like small arboreal marsupial didelophids, like Caluromys, and small arboreal placental tree shrews, like the extant Ptilocercus and Tupaia. Two distinct specimens, both given the genus name Hapalodectes (Fig. 1), are among these basal placental taxa in the large reptile tree (LRT, 1378 taxa).

The slightly smaller
IVPP V5235 specimen attributed to Hapalodectes (Ting and Li 1987) nests at the base of the primate clade. It had already taken on the appearance of a little basal lemur or adapid (Fig. 1).

Figure 1. Two Hapalodectes specimens. The smaller one nests at the base of the Primates. The larger one nests as the base of the anagalid-tenrec-odontocete clade.

Figure 1. Two Hapalodectes specimens. The smaller one nests at the base of the Primates. The larger one nests as the base of the anagalid-tenrec-odontocete clade.

The slightly larger
IVPP V12385 specimen attributed to Hapalodectes (Ting et al. 2004; Fig. 1) nests at the base of the anagalid-tenrec-odontocete clade and it had already taken on the appearance of a little anagalid or elephant shrew.

Other than size, the differences are subtle:

  1. The basal primate has a postorbital ring. The basal anagalid does not.
  2. The basal primate has three upper molars. The basal anagalid has four.
  3. The basal primate cranium has no crest. The basal anagalid has a nuchal and parasagittal crest.
  4. The basal primate anchors the squamosal further back, with a smaller ectotympanic (middle ear container bones below the cranium). The basal anagalid anchors the squamosal further forward, with a larger ectotympanic (for better hearing).

Hapalodectes hetangensis (Ting and Li 1987; 4.5cm skull length; Paleocene, 55 mya; IVPP V 5235) This skull was originally wrongly applied to the Mesonychidae, but here nests at the base of the primates, including Notharctus.  Note the transverse premaxilla, the large canine, and the encircled orbits rotated anteriorly.

?Hapalodectes ?hetangensis (Ting et al. 2004; 7 cm skull length; Early Eocene 50 mya; IVPP V 12385) was originally considered a tiny mesonychid. This species nests at the base of the anagale-tenrec-odontocete clade along with Maelestes, between Ptilocercus and Onychodectes. The large nuchal crest is a key trait found in later taxa. The premaxilla is largely missing, but likely was transverse in orientation.

Figure 2. Ptilocercus (pen-tailed tree shrew) compared to Caluromys (wooly-opossum) young juvenile from Flores, Abdala and Giannini 2010.

Figure 2. Ptilocercus (pen-tailed tree shrew) compared to Caluromys (wooly-opossum) young juvenile from Flores, Abdala and Giannini 2010.

References
Ting S and Li C 1987. The skull of Hapalodectes (?Acreodi, Mammalia), with notes on some Chinese Paleocene mesonychids.
Ting SY, Wang Y, Schiebout JA, Koch PL, Clyde WC, Bowen GJ and Wang Y 2004. New Early Eocene mammalian fossils from the Hengyang Basin, Hunan China. Bulletin of Carnegie Museum of Natural History 36: 291-301.

wiki/Hapalodectes

The prehensile hand and foot of Caluromys

Figure 1. Pteropus and Caluromys compared in vivo and three views of their skulls. Caluromys is in the ancestry of bats and shows where they inherited their inverted posture.

Figure 1. Pteropus and Caluromys compared in vivo and three views of their skulls. Caluromys is in the ancestry of bats and shows where they inherited their inverted posture.

I could not find
and still cannot find a complete skeleton for Caluromys (Fig. 1), the transitional marsupial leading to placentals. Argot 2001 published images of the hand in vivo. Argot 2002 published images of the foot in vivo and as an incomplete set of bones (Fig. 2). I matched those bones to the foot, still wishing I had all the bones, as in an X-ray.

Figure 1. Caluromys hand and foot from Argot 2002 compared to Didelphis and repaired here to match.

Figure 2. Caluromys hand and foot from Argot 2001, 2002 compared to the pes of Didelphis. The manus and pes of primates, tree shrews (in Glires) and basal arboreal Carnivorans all arise from Caluromys. These demonstrate the early appearance of the prehensile/opposable big toe and thumb, derived from the semi-opposable big toe of Didelphis, the Virginia opossum and even more so in Caluromys. 

The prehensile manus and pes of Caluromys
is primitive for the Eutheria (= Placentalia). From these arise the wings of bats, the flippers of whales, the hooves of horses as well as the fingers I just used to type this sentence.

References
Argot C 2001. Functional-Adaptive Anatomy of the Forelimb in the Didelphidae, and the Paleobiology of the Paleocene Marsupials Mayulestes ferox and Pucadelphys andinus. Journal of Morphology 247:51–79.
Argot C 2002. Functional-adaptive analysis of the hindlimb anatomy of extant marsupials and the paleobiology of the Paleocene marsupials Mayulestes ferox and
Pucadelphys andinus. Journal of Morphology 253:76–108.

Basal placentals illustrated in phylogenetic order

Eutherian (= placental) mammals
are divided into clades like Primates, Ungulata, Carnivora, etc. Known basal taxa for each of these clades are related to one another in a ladder-like fashion, each one nesting at the base of a bushy clade in the large reptile tree (LRT, 1366 taxa).

Today,
an illustration of skeletons (Fig. 1) and skulls (Fig. 2) in phylogenetic order documents the minor changes (microevolution) between basal taxa that nest at the bases of several increasingly derived placental clades.

FIgure 1. Skeletons of taxa that nest at the bases of several major placental clades, divided between Cretaceous and Paleocene taxa.

FIgure 1. Skeletons of taxa that nest at the bases of several major placental clades, divided between Cretaceous and Paleocene taxa, divided by four different scales. Basal taxa are several degrees of magnitude smaller.

The following placental skulls are not to scale
yet continue to demonstrate the minor changes (microevolution) that occur at the bases of several major placental clades. For instance, Chriacus is basal to bats, while Maelestes is basal to odontocete whales. It is difficult, if not impossible, to determine such future developments in these basal taxa without the benefit of a wide gamut analysis, like the LRT.

Figure 2. A selection of placental skulls in phylogenetic order and divided into Cretaceous and Paleocene taxa.

Figure 2. A selection of placental skulls in phylogenetic order and divided into Cretaceous and Paleocene taxa.

The lesson for today:
Sometimes quantity, without firsthand observation, is needed to put together the ‘Big Picture’ before one is able to pick apart the details that each specific specimen reveals during firsthand study. Traditionally paleontologists have been putting the latter ahead of the former by (too often) excluding pertinent taxa revealed and documented by the more generalized and wide gamut phylogenetic analysis provided by the LRT. Like Yin and Yang, both must be considered. ‘Avoid taxon omission‘ is the single most important rule when constructing a cladogram of interrelationships.

References
See ReptileEvolution.com and links therein.

Mesozoic mammals: Two views

Smith 2011 reported,
at the beginning of the Eocene, 55mya, “the diversity of certain mammal groups exploded.” These modern mammals”, according to Smith, ‘ consist of rodents, lagomorphs, perissodactyls, artiodactyls, cetaceans, primates, carnivorans and bats. Although these eight groups represent 83% of the extant mammal species diversity, their ancestors are still unknown. A short overview of the knowledge and recent progress on this research is here presented on the basis of Belgian studies and expeditions, especially in India and China.’

Contra the claims of Smith 2011
in the large reptile tree (LRT, 1354 taxa, subsets Figs. 2–4) prototherians are known from the late Triassic (Fig. 1). Both metatherians and eutherians are known from the Middle Jurassic. Many non-mammal cynodonts survived throughout the Mesozoic. In addition, the ancestors of every included taxon are known back to Devonian tetrapods.

Noteworthy facts after an LRT review (Fig. 1):

  1. All known and tested Mesozoic mammals (Fig. 1) are either small arboreal taxa or small burrowing taxa (out of sight of marauding theropods).
  2. All Mesozoic monotremes are more primitive than Ornithorhynchus and Tachyglossus (both extant).
  3. All Mesozoic marsupials are more primitive than or include Vintana (Late Cretaceous).
  4. All Mesozoic placentals are more primitive than Onychodectes (Paleocene).
Figure 1. Select basal cynodonts and mammals set chronologically. The divergence times for placentals (Eutheria), marsupials (Metatheria) and monotremes (Mammalia) are estimated here.

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

Given those parameters
we are able to rethink which mammals were coeval with dinosaurs back on phylogenetic bracketing (= if derived taxa are present, primitive taxa must have been present, too).

Smith reports, “The earliest known mammals are about as old as the earliest dinosaurs and appeared in the fossil record during the late Trias around two hundred and twenty million years ago with genera such as Sinoconodon (pre-mammal in the LRT), Morganucodon (basal therian in the LRT) and Hadrocodium (basal therian in the LRT). However, the earliest placental mammals (Eutheria) were not known before the Early Cretaceous. Eomaia scansoria (not eutherian in the LRT) from the Barremian of Liaoning Province, China is the oldest definite placental and is dated from a hundred and thirty million years ago.”

Mesozoic Prototherians

  1. All included fossil taxa are Mesozoic. Two others are extant (Fig. 2).
Figure 2. Mesozoic prototherians + Megazostrodon, the last common ancestor of all mammals. Only two taxa (gray) are post-Cretaceous.

Figure 2. Mesozoic prototherians + Megazostrodon, the last common ancestor of all mammals. Only two taxa (gray) are post-Cretaceous.

Mesozoic Metatherians (Marsupials)

  1. Derived Vincelestes is Early Cretaceous, which means Monodelphis and Chironectes were present in the Jurassic.
  2. Derived Didelphodon is Late Cretaceous, which means sisters to Thylacinus through Borhyaena were also present in the Mesozoic.
  3. Derived Vintana is Late Cretaceous, which means sisters to herbivorous marsupials were also present in the Mesozoic.
Figure 3. Mesozoic metatherians (in black), later taxa in gray. Whenever derived taxa are present in the Mesozoic (up to the Late Cretaceous) then ancestral taxa, or their sisters, were also present in the Mesozoic. Didelphis is extant, but probably unchanged since the Late Jurassic/Early Cretaceous.

Figure 3. Mesozoic metatherians (in black), later taxa in gray. Whenever derived taxa are present in the Mesozoic (up to the Late Cretaceous) then ancestral taxa, or their sisters, were also present in the Mesozoic. Didelphis is extant, but probably unchanged since the Late Jurassic/Early Cretaceous.

Mesozoic Eutherians (= Placentals)

  1. Rarely are placental mammals identified from the Mesozoic, because many are not considered placentals.
  2. Placentals (in the LRT) are remarkably rare in the Mesozoic, but sprinkled throughout the cladogram, such that all taxa more primitive than the most derived Mesozoic taxon (Maelestes and derived members of the clade Glires, Fig. 4, at present a number of multituberculates) must have had Mesozoic sisters (Carnivora, Volitantia, basal Glires). 
Figure 4. Mesozoic euthrerians (placentals, in black). Later taxa in light gray. All taxa more primitive than Mesozoic taxa were likely also present in the Jurassic and Cretaceous. None appear after Onychodectes. Madagascar separated from Africa 165-135 mya, deep into the Cretaceous with a population of tenrecs attached. No rafting was necessary. 

Figure 4. Mesozoic euthrerians (placentals, in black). Later taxa in light gray. All taxa more primitive than Mesozoic taxa were likely also present in the Jurassic and Cretaceous. None appear after Onychodectes. Madagascar separated from Africa 165-135 mya, deep into the Cretaceous with a population of tenrecs attached. No rafting was necessary.

The above represents what a robust cladogram is capable of,
helping workers determine the likelihood of certain clades appearing in certain strata, before their discovery therein, based on their genesis, not their widest radiation or eventual reduction and extinction. In other words, we might expect sisters to basal primates, like adapids and lemurs, to be present in the Mesozoic, but not sisters to apes and hominids. We should expect sisters to all tree shrews and rodents to be recovered in Mesozoic strata. We should expect to see sisters to Caluromys, Vulpavus and other small arboreal therians/carnivorans in Mesozoic strata, but not cat, dog and bear sisters.

References
Smith T 2011. Contribution of Asia to the evolution and paleobiogeography of the earliest modern mammals. Bulletin des séances- Académie royale des sciences d’outre-mer. Meded. Zitt. K. Acad. Overzeese Wet.57: 293-305