Acostasaurus enters the LRT

Pérez and Noé 2017 described
a near complete 3D skull, a complete hindlimb and several vertebrae of a eusauropterygian, Acostasaurus (Fig. 1), they considered it a 4-5m long, snort-snouted pliosaur, one of many ‘pliosaurs’ found in Barremian (Early Cretaceous) Columbia. 

Figure 1. Acostasaurus skull from Pérez and Noé 2017, colors added.

Figure 1. Acostasaurus skull from Pérez and Noé 2017, colors added.

Some of those purported Columbian ‘pliosaurs’
turned out to be giant sisters to more basal eusauropterygians in the large reptile tree (LRT, 1430 taxa). You might remember (here) the giant Sachicasaurus nested with Nothosaurus and (here) the very large Bobosaurus nested with the smaller Corosaurus.

In the LRT
Acostasaurus nests with Anningsaura (Fig. 2) apart from the pliosaurs in the LRT.

Figure 6. Anningasaura colorized from an old engraving. No other aquatic taxon has such bizarrely curved teeth. This taxon is closely related to Hauffiosaurus.

Figure 6. Anningasaura colorized from an old engraving. No other aquatic taxon has such bizarrely curved teeth. This taxon is closely related to Hauffiosaurus.

The authors compared Acostasaurus
with Simolestes a taxon not yet added to the LRT. Look for it soon.

Figure 4. Subset of the LRT focusing on Eusauropterygians (pachypleurosaurs, nothosaurs, plesiosaurs and kin).

Figure 4. Subset of the LRT focusing on Eusauropterygians (pachypleurosaurs, nothosaurs, plesiosaurs and kin).


References
Gómez Pérez M and Noè LF 2017. Cranial anatomy of a new pliosaurid Acostasaurus pavachoquensis from the Lower Cretaceous of Colombia, South America. Palaeontographica Abteilung A. 310 (1–2): 5–42. doi:10.1127/pala/2017/0068.

wiki/Acostasaurus

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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

SVP 2018: Placentals in the Cretaceous

Halliday et al. 2018
wonder about “the traditional lack of Cretaceous placental fossils when results from diverse dating analyses favor a Cretaceous origin of Placentalia.”

Unfortunately
they use an outdated cladogram that includes the following invalid clades (superorders) that Halliday et al. surmise should be present in Cretaceous sediments:

  1. Atlantogenata = Afrotheria + Xenarthra (elephants and anteaters in one clade?)
  2. Laurasiatheria = shrews, pangolins, bats, whales, carnivorans and ungulates (whales and bats in the same clade?)
  3. Euarchontoglires = rodents, lagomorphs, tree shrews, colugos and primates (lacking only carnivores, these are basal eutherians

Together these three clades
comprise the entirety of extant Eutheria (placental mammals). All of the above clades are extant. Where are the extinct clades, like Multituberculata?

By contrast,
the large reptile tree  (LRT, 1313 taxa) recovers Middle and Late Jurassic placentals (multituberculate rodents) along with several Early Cretaceous taxa, like the pangolin ancestor, Zhangheotherium (Fig. 1). So “the traditional lack of Cretaceous placental fossils” has been updated in the LRT.

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.

Halliday et al. conclude: “The lack of definitive Cretaceous placental mammals may therefore be explained by high predicted morphological similarity among stem and basal crown eutherians, providing an avenue for partially reconciling the fossil record and molecular divergence estimates in Placentalia.”

No.
Taxon exclusion has given Halliday et al. an outdated tree topology. There is plenty of evidence for Mesozoic placentals in the LRT. Adding taxa provides every included taxon new opportunities to nest more parsimoniously. A good starter list can be found here (LRT subset Fig. 2). Many taxa from this list are candidates for discovery in the Mesozoic based on the discovery of multituberculates in the Mesozoic.

Figure 3. Subset of the LRT, focusing on basal Eutheria and Heterocephalus. Aqua taxa are arboreal. Tan taxa are terrestrial. Blue taxa are aquatic.

Figure 3. Subset of the LRT, focusing on basal Eutheria and Heterocephalus. Aqua taxa are arboreal. Tan taxa are terrestrial. Blue taxa are aquatic.

References
Halliday TJ et al. (5 co-authors) 2018. Delayed increase in morphological rates of evolution after the origin of the placental mammal crown group. SVP abstracts.

Mammal taxa: origin times

A few days ago, we looked at a revised and expanded cladogram of the Mammalia based on skeletal traits (distinct from and contra to a cladogram based on DNA). Today we add chronology to the cladogram to indicate the first appearance of various mammals and estimate the origin of the various clades (Fig. 1).

Note that derived taxa
that chronologically precede more primitive taxa indicate that primitive taxa had their genesis and radiation earlier than the first appearance of fossil specimens, which always represent rare findings usually during wide radiations that increase the chance the specimen will fossilize in the past and be found in the present day.

Looking at time of mammal taxa origin categories:

Figure 1. Cladogram with time notes for the Mammalia (subset of the LRT).

Figure 1. Cladogram with time notes for the Mammalia (subset of the LRT).

Some notes:

  1. Both prototheres and basal therians were present (and probably widespread) in the Late Triassic.
  2. Derived prototheres appear in the Late Triassic, suggesting an earlier (Middle Triassic?) origin for Mammalia and an earlier (Middle Triassic?) split between Prototheria and Theria.
  3. Both fossorial metatherians and basal arboreal eutherians were present (and probably widespread) in the Late Jurassic. These were small taxa, out of the gaze of ruling dinosaurs.
  4. Large derived eutherians eolved immediately following the K-T boundary in the Paleocene and radiated throughout the Tertiary.
  5. A large fraction of prototherians, metatherians and eutherians are known only from extant taxa, some of which are rare and restricted, not widespread.
  6. Multituberculates and kin are derived placentals close to rodents by homology, not convergence.

 

‘The Dawn of Mammals’ YouTube video illuminates systematic problems

Sorry, looks like video was yanked off of YouTube.

Part of this YouTube video (see below, click to view)
pits DNA paleontolgist, Dr. Olaf Bininda-Emonds (U. Oldenburg), against bone trait paleontologist, Dr. John Wible (Carnegie Museum of Natural History) in their common and contrasting search for basal placental mammals. Both realize that DNA cladograms do not replicate bone cladograms and DNA cannot be utilized with ancient fossils.

Dr. Bininda-Emonds, used molecular clocks
in living taxa to hypothetically split marsupials from placentals about 160 mya ago (Late Jurassic).

By contrast, Dr. Wible reports (28:53),
“Our study supported the traditional view that there were no fossils living during the Cretaceous [that] were members of the placental group itself. There were only ancestors of the placentals living.” (unscripted verbatim)

The impulse for this argument
came from the discovery of Maelestes (Wible et al. 2007a,b; 28:30 on the video) from the Late Cretaceous (75 mya). Dr. Wible’s paper nested Maelestes with the pre-placental, Asiorcytes, another tree-shrew-like mammal from the Late Cretaceous. By contrast, the LRT, nests Maelestes unequivocally at the base of the tenrec/odontocete clade, well within the placental clade (Fig. 2), as we learned earlier here.

The large reptile tree
 (subset in Fig. 2) nests the first known placental mammals at the 160 mya mark, matching the DNA predictions of Dr. Bininda-Emonds et al. A long list of taxa, including Maelestes, nest in the Jurassic and Cretaceous, contra Wible et al. Only more complete taxa are tested in the LRT and dental traits are not emphasized.

Figure 2. Mesozoic time line showing the first appearances of several fossil mammals and the clades they belong to. Many, if not most of the listed taxa are late survivors of earlier radiations, sometimes much earlier radiations. Monodelphis and Didelphis are extant animals that originated in the Early Jurassic at the latest. Note also the large gaps over tens of millions of years, highlighting the rarity of fossil bearing locales.

Figure 2. Mesozoic time line showing the first appearances of several fossil mammals and the clades they belong to. Many, if not most of the listed taxa are late survivors of earlier radiations, sometimes much earlier radiations. Monodelphis and Didelphis are extant animals that originated in the Early Jurassic at the latest. Note also the large gaps over tens of millions of years, highlighting the rarity of fossil bearing locales.

In the video Dr. Wible says, “Many modern groups, according to the molecular clock analysis, actually are, they should be, present in the Cretaceous fossil record. We can’t find them.” Actually Dr. Wible already found them, but does not recognize them for what they are. That’s a common problem in paleontology, largely due to taxon exclusion, that we’ve seen before here, here, here and here. And in dozens of other mislabeled clades, like multituberculates.

The Bininda-Edmonds et al. paper reports,
“Here we construct, date and analyse a species-level phylogeny of nearly all extant Mammalia to bring a new perspective to this question. Our analyses of how extant lineages accumulated through time show that net per-lineage diversification rates barely changed across the Cretaceous/Tertiary boundary. Instead, these rates spiked significantly with the origins of the currently recognized placental superorders and orders approximately 93 million years ago, before falling and remaining low until accelerating again throughout the Eocene and Oligocene epochs. Our results show that the phylogenetic ‘fuses’ leading to the explosion of extant placental orders are not only very much longer than suspected previously, but also challenge the hypothesis that the end-Cretaceous mass extinction event had a major, direct influence on the diversification of today’s mammals.”

The LRT agrees with the timing indicated by the DNA analysis
Placentals are indeed found in the LRT Cretaceous and Jurassic fossil record (Fig. 2). They were not recognized by traditional workers using smaller taxon lists, for what they were. The LRT minimizes taxon exclusion and so solves many paleo problems with an unbiased and wide gamut approach currently unmatched in the paleo literature. Extant birds have a similar deep time record based on a few recent finds.

Perhaps overlooked
there are currently large gaps spanning tens of millions of years, highlighting the rarity of fossil bearing locales. All Mesozoic mammals are rare.

The DNA tree
of the Bininda-Emonds team correctly splits monotremes from therians, but incorrectly nests ‘Afrotherians‘ with Xenarthrans at the base of all mammals followed by moles + shrews, bats + carnivores + hoofed mammals + whales, followed by primates and rodents. As anyone can see, this is a very mixed up order, placing small arboreal taxa in derived positions and stiff-backed elephants and in in basal nodes. This DNA analysis is not validated by the LRT.

To its credit, basal mammals in the LRT
greatly resemble their marsupial ancestors. Then derived mammals become generally larger, with derived tooth patterns, stiffer dorsal/lumbar areas and longer pregnancies with more developed (precocious) young.

Given three cladograms of placental relationships,
none of them identical, how does one choose which one is more accurate? Here’s my suggestion: look at each sister at each node and see where you best find a gradual accumulation of derived traits, without exception. And look at outgroups leading to basal members of the in group.

Some readers are still having a hard time realizing
that someone without direct access to fossils and without a PhD is able to recover a more highly resolved cladogram that features gradual changes between every set of sister taxa than trees published over the last ten years in the academic literature. I agree. This should not be taking place. This is not what I expected to find when I started this 7-year project. One tends to trust authority. It’s been an eye-opening journey. In nearly all tested studies overlooking relevant taxa continues to be the number one shortcoming. The LRT minimizes that issue. The number two problem is blind faith in DNA results. The number three problem is an apparent refusal to examine phylogenetic results to weed out mismatched recovered sister taxa.

The video spends also some time with Zhangheotherium,
which we looked at earlier here and here. The interviewed workers talk about the ankle spur, but as a venom injector, as in the duckbill, Ornithorhynchus, not as a membrane frame, like a calcar bone, as in bats.

The video considers Repenomamus a large Early Cretaceous mammal
but the LRT nests Repenomamus as a late-surviving synapsid pre-mammal, derived from a sister to Pachygenelus, as we learned earlier here.

PS. As touched on earlier,
many basal arboreal mammals were experimenting with gliding (e.g. Volaticotherium and Maiopatagaium), but only one clade, bats, experimented with flapping. This was, perhaps not coincidentally, during the Middle to Late Jurassic (Oxordian, 160 mya). Remember, these gliding membranes were all extensions of the infant nursery membrane found in colugos and other volatantians, not far from the basalmost placental, Monodelphis.

References
Bininda-Emonds ORP, et al., (9 co-authors) 2007. The delayed rise of present-day mammals. Nature 446(7135):507-512.
Wible JR, Rougier GW, Novacel MJ and Asher RJ 2007a. The eutherian mammal Maelestes gobiensis from the Late Cretaceous of Mongolia and the phylogeny of Cretaceous Eutheria. Bulletin of the American Museum of Natural History 327:1–123.
Wible JR, Rougier GW, Novacek MJ and Asher RJ 2007b. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary.” Nature, 447: 1003-1006.

 

What is a limpkin? (genus: Aramus)

Figure 1. The limpkin (Aramus guarauna) is a basal member of the x family.

Figure 1. The limpkin (Aramus guarauna) is a long-legged, wading basal member of the x family.

Aramus guarauna (Linneaus 1766) is the extant limpkin. It is often considerd transitional between rails and cranes. In the large reptile tree (1121 taxa) the limpkin nests basal to seagulls and hummingbirds, plovers and crowned cranes, common cranes and stilts, terns and loons, kingfishers and jabirus, murres and penguins.

Figure 1. Skeleton of the limp kin (Aramus), traditionally nests within the crane and rail order Gruiformes.

Figure 2. Skeleton of the limpkin (Aramus), traditionally nests within the crane and rail order Gruiformes. In the LRT rails are not closely related, so Gruiformes should no longer include rails.

Extant limpkins eat snails.
Primitive limpkins like Aramournis  probably had a more diverse diet. It is known from a distal tarsus.

Traditional rails
like the corn crake (Crex) and the coot (Fulica) are much more basal birds that give rise to chickens, sparrows and parrots. Adding Rallus, the Virginia rail, to the LRT nests it between Aramus and the rest of the clade, which, phylogenetically makes hummingbirds, terns and penguins variations on the rail theme and Rallus at least a Middle Cretaceous taxon radiation.

Figure 4. Virginia rail alongside the rail clade in the LRT.

Figure 4. Virginia rail alongside the rail clade in the LRT.

Congeneric specimens of Aramus
are found in the Miocene, but more derived penguins are found in the Paleocene, pointing to a mid-Cretaceous radiation of this clade.

Limpkins are derived from Cretaceous sisters to
hamerkops (Scopus) and stone curlews (Burhinus), both long-legged taxa. By the evidence shown in the crown bird subset of the LRT (Fig. 4), long legs, like those shown by Aramus, the limpkin, are basal traits. The retention of hatchling short legs occurred several times by convergence, sometimes during the Cretaceous. See the earlier post on post K-T non-arboreal birds. 

Figure 4. Subset of the LRT focusing on the crown bird clade. Brown taxa are all long-legged. Neotony produces the smaller, shorter-legged, arboreal taxa.

Figure 4. Subset of the LRT focusing on the crown bird clade. Brown taxa are all long-legged. Neotony produces the smaller, shorter-legged, arboreal taxa.

References
Linneaus C von 1766. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. pp. 1–532. Holmiæ. (Salvius)

wiki/Aramus_limpkin

Bird, pterosaur, dinosaur simplified chronology

Following the earlier post on non-arboreal post K-T boundary birds…

…this one pretty much speaks for itself.
Here (Fig. 1) is a chronology, very much simplified, of birds, pterosaurs and dinosaurs according to the LRT.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades based on taxa in the LRT.

If you’re curious about any of the taxa,
in the chronology, simply use Keywords to locate them.