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they use an outdated cladogram that includes the following invalid clades (superorders) that Halliday et al.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 an outdated tree topology.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.

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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

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.

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.

Early Cretaceous stem chameleon/horned lizard

Unnamed stem chameleon (Daza et al. 2016; Early Cretaceous, 1.2cm in length; JZC Bu154; Fig. 1) is a tiny neonate preserved in amber. It also nests basal to horned lizards like Phrynosoma, in the large reptile tree (LRT, 1089 taxa). Note the long, straight hyoid forming the base of the shooting tongue. The split fingers and toes of extant chameleons had not yet developed in this taxon. Found in amber, this newborn lived in a coniferous forest.

Figure 1. The Early Cretaceous stem chameleon/horned lizard found amber. Snout to vent length is less than 11 mm. Much smaller than a human thumbnail.

Figure 1. The Early Cretaceous stem chameleon/horned lizard found amber. Snout to vent length is less than 11 mm. Much smaller than a human thumbnail. Insitu fossil from Daza et al. 2016,  colorized and reconstructed here. At a standard 72 dpi screen resolution, this specimen is shown 10x actual size.

This specimen further cements
the interrelationship of arboreal chameleons and their terrestrial sisters, the horned lizard we looked at earlier with Trioceros and Phyrnosoma in blue of this cladogram (Fig. 2) subset of the LRT.

Figure 3. Subset of the LRT focusing on the neonate stem chameleon/horned lizard.

Figure 2. Subset of the LRT focusing on the neonate stem chameleon/horned lizard.

Figure 6. Phyronosoma, the horned lizard of North America.

Figure 3. Phyronosoma, the horned lizard of North America.

Figure 2. Trioceros jacksonii overall. Size is 12 inches (30 cm) from tip to tip.

Figure 4. Trioceros jacksonii overall. Size is 12 inches (30 cm) from tip to tip.

References
Daza JD et al. 2016. Mid-Cretaceous amber fossils illuminate the past diversity of tropical lizards. Sci. Adv. 2016; 2 : e1501080 4 March 2016

New flightless and giant nyctosaurs: Alcione and Barbaridactylus

Scale bar problems
and a lack of reconstructions in the original paper are issues here.

Longrich, Martill and Andres 2018
bring us news of “a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. [This] dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families —Pteranodontidae, Nyctosauridae, and Azhdarchidae — persisted into the late Maastrichtian. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.”

The authors summary starts off with an invalid statement:
“Pterosaurs were winged cousins of the dinosaurs.”  That was invalidated by Peters 2000, 2007 and ignored every since. We looked at that problem earlier here, here and here in a 3-part series testing all candidates. It’s time to realize that no one will ever find pterosaur kin among the dinos. They’ve already been clearly identified among the lepidosaurs.

The authors failed to include the Maastrictian tupuxuarid
found in southern Texas (Fig. 1; TMM 42489-2) and did not consider the Maastrichtian footprints discovered in 1954 and reexamined in 2018 that include two ctenochasmatids we will look at tomorrow.

TMM 42489-2, the tall crested Latest Cretaceous large rostrum and mandible. It's a close match to that of Tupuxuara, otherwise known only from Early Cretaceous South American strata.

Figure 1. TMM 42489-2, the tall crested Latest Cretaceous large rostrum and mandible. It’s a close match to that of Tupuxuara, otherwise known only from Early Cretaceous South American strata.

Alcione elainus gen. et sp. nov.
The new 1.5x larger nyctosaurid, Alcione elainus, known from disassociated bones including a shorter radius + ulna, a shorter metacarpal 4, a larger femur, and a tiny sternal complex (identified as a ‘sternum’ in the text) only 40 percent the size of a standard nyctosaur sternal complex (if the scale bars are correct). When placed on a reconstruction of a more complete Nyctosaurus (UNSM 93000; Fig. 2), scaled to the humerus, the result produces a likely flightless nyctosaur. Strangely, the authors called this a “small nyctosaur” even though it is half again larger than UNSM 93000. The authors mislabeled the shorter, straighter scapula as a coracoid, and vice versa.

Figure 2. GIF movie of Nyctosaurus and Alcione showing a likely flightless nyctosaur based on the parts preserved.

Figure 2. GIF movie of Nyctosaurus and Alcione showing a likely flightless nyctosaur based on the parts preserved. Three frames change every 5 seconds. The sternum is tiny (assuming the scale bars are correct), the metacarpus and antebrachium are short and the femur is long.

They did not mention the possibility of flightlessness.
They did report, “The abbreviated distal wing elements in Alcione indicate a specialized flight style. The short, robust proportions suggest reduced wingspan and increased wing loading, implying distinct flight mechanics and an ecological shift. Short wings would increase lift-induced drag at low speeds, but reduced wing areas would decrease parasite drag at high speeds, suggesting that Alcione may have been adapted for relatively fast flapping flight compared to other nyctosaurids. Alternatively, reductions in wingspan might represent an adaptation to underwater feeding, i.e., plunge diving of the sort practiced by gannets, tropicbirds, and kingfishers, where smaller wings would reduce drag underwater.”

Not sure why they mentioned
‘distal wing elements’ here. They did not list or discuss distal wing elements elsewhere. Perhaps they meant proximal.

The reconstructed mandible of Alcione
is narrower than the rostrum in UNSM 93000.

Based on the vestigial fingers of UNSM 93000
and the short metacarpus of the new specimen, Alcione might have been the first pterosaur to walk on metacarpal 4, albeit at the very end of the reign of pterosaurs.

Other flightless pterosaurs include:
the basal azhdarchid form the Solnhofen, Jme-Sos 2428 and the Late Jurassic anurognathid PIN 2585/4 from the Sordes slab. They demonstrate that the distal wing elements reduce first. Thus the reconstruction, based on nyctosaur patterns restores a wing that was not volant.

Longrich, Martill and Andres did find a giant nyctosaur
which they named Barbaridactylus grandis based on a large humerus (Fig. 3). The humerus of the more complete UNSM 93000 specimen is 9.5 cm. By comparison the humerus in Barbaridactylus is 22.5 cm. I’m going to trust the text comment that the ulna + radius are 1.3x longer than the humerus. The scale bars indicate about half that length. Similar problem possible in the scapula/coracoid, according to the nyctosaur bauplan.

Figure 3. Barbaridactylus, a giant nyctosaurid. If the wing was like UNSM 93000, then it could fly. If the wing was like Alcione, then it could not. The scale bars did not match the text description on the ulna + radius, so both sizes are shown.

Figure 3. Barbaridactylus, a giant nyctosaurid. If the wing was like UNSM 93000, then it could fly. If the wing was like Alcione, then it could not. The scale bars did not match the text description on the ulna + radius, so both sizes are shown. Sometimes you have to be prepared for the occasional mistake in a published paper.

Other giant nyctosaurs
Earlier and here we noted giant nyctosaurs were flying over the Niobrara Sea (midwest North America) based on a large wing finger with unfused extensor tendon process (YPM 2501) and a large nyctosaur pelvis (KUVP 993; misinterpreted by Bennett (1991, 1992) as belonging to a female Pteranodon). 

No reconstructions were provided
by Longrich, Martill and Andres 2018. Reconstructions and a nyctosaur blueprint might have helped these paleontologists with firsthand access to the specimens discover the issues they missed.

It’s good to know
more pterosaurs made it to the latest Cretaceous.

References
Bennett SC 1991. Morphology of the Late Cretaceous Pterosaur Pteranodon and Systematics of the Pterodactyloidea. [Volumes I & II]. Ph.D. thesis, University of Kansas, University Microfilms International/ProQuest.
Bennett SC 1992.
 Sexual dimorphism of Pteranodon and other pterosaurs, with comments on cranial crests. Journal of Vertebrate Paleontology 12: 422–434.
Longrich NR, Martill DM, Andres B 2018.
Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary. PLoS Biol 16(3): e2001663. https://doi.org/10.1371/journal.pbio.2001663
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2007. 
The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.

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