Laosuchus naga enters the LRT

The question today is:
what are chroniosuchians? Are they reptiles or not? Arbez, Sidor and Steyer 2018 say: ‘not’ (Fig. 1). Here that mistake is due to tradition and taxon exclusion, based on their cherry-picked outgroups. Heretically. chroniosuchians are amphibian-like reptiles.

Figure 1. Cladogram from xx 2018 with Laosuchus nesting with chroniosuchians in the absence of Solenodonsaurus.

Figure 1. Cladogram from Arbez, Sidor and Steyer 2018 with Laosuchus nesting with chroniosuchians in the absence of Solenodonsaurus.

Arbez, Sidor and Steyer report from their abstract:
“Chroniosuchians were a clade of non-amniotic tetrapods known from the Guadalupian (middle Permian) to Late Triassic, mainly from Russia and China.” Asaphestera is the proximal outgroup followed by Limnoscelis, Seymouria, Gephyrostegus and other taxa.

By contrast and using more outgroup taxa
the large reptile tree (LRT 1301 taxa, subset Fig. 2) nests chroniosuchians within the base of the archosauromorph branch of reptiles. When more taxa are included in the LRT, Limnoscelis and Gephyrostegus nest as reptiles (= amniotes) while Asaphestra and Seymouria nest as unrelated traditional microsaur lepospondyls and seymouriamorphs respectively.

Figure 1. Laosuchus nests with untested Solenodonsaurus rather than the extra fenestrated chroniosuchians.

Figure 2. Laosuchus nests with untested Solenodonsaurus rather than the extra fenestrated chroniosuchians. Note that all these reptiles precede the traditional first dichotomy in amniote cladograms: the splitting of synapsida from the other amniotes.

Arbez, Sidor and Steyer 2018 introduce a new taxon,
Laosuchus naga (Fig 3), a long-snouted chroniosuchian without an antorbital fenestra. The authors did not include Solenodonsaurus (Fig. 3), which nests with Laosuchus as basal chroniosuchians in the LRT (Fig. 2). Both lack an antorbital fenestra, distinct from Chrionosaurus and Chroniosuchus.  

Figure 2. Laosuchus compared to Bystrowiella and Solenodonsaurus to scale (presuming that the scale bar for Laosuchus is 10cm.)

Figure 2. Laosuchus compared to Bystrowiella and Solenodonsaurus to scale (presuming that the scale bar for Laosuchus is 10cm.) Distinct from many reptiles, the lacrimal no longer contacts the orbit in this taxon and other chroniosuchians, but continues to contact the naris.

Laosuchus naga traits include:

  1. an extremely reduced pineal foramen
  2. absence of palatal dentition
  3. well-developed transverse flange of the pterygoid that contacts the maxilla
  4. internal crest on and above the dorsal side the palate
  5. otic notch closed by the tabular horn and the posterior part of the squamosal, forming a continuous wall
  6. thin and high ventromedial ridge on parasphenoid.
Figure 4. Solenodonsaurus skull in situ and reconstructed. That brown bone on top of the frontal/parietal suture is a displaced lacrimal that nicely fills the gap in the reconstruction.

Figure 4. Solenodonsaurus skull in situ and reconstructed using DGS. That brown bone on top of the frontal/parietal suture are displaced pieces of the lacrimal that nicely fills the gap in the reconstruction. A few bones are always ignored. I’m just trying to get the basics here.

Something I learned while reexamining Solenodonsaurus
The displaced bone atop the skull is actually part of the broken lacrimal. The quadratojugal is displaced on the posterior mandible. The prefrontal is broken but not very displaced. The posterior jugal is broken into several pieces. Using DGS allows one to cut and paste and fit these puzzle pieces back into the missing parts of the skeleton where they belong. If they don’t fit, they don’t belong, but they never fit perfectly. It’s like putting Humpty Dumpty together again. There are always a few pieces left over.

References
Arbez T, Sidor CA and Steyer J-S 2018. Laosuchus naga gen. et sp. nov., a new chroniosuchian from South-East Asia (Laos) with internal structures revealed by micro-CT scan and discussion of its palaeobiology. Journal of Systematic Palaeontology DOI: 10.1080/14772019.2018.1504827

http://zoobank.org/urn:lsid:zoobank.org:pub:11D07FA3-0F4C-4EF9-A416-E8E6BE76C970

 

Advertisements

Wachtlerosaurus: a thalattosaur, not an archosaur

Perner 2018
introduces Wachtlerosaurus ladinicus (Fig. 1, 2), a tiny disarticulated reptile from the Middle Triassic Dolomites of northern Italy. Perner considers the specimen an archosaur. In the large reptile tree (LRT, 1264 taxa) the specimen nests with thalattosaurs and Thalattosaurus in particular. The purported antorbital fenestra is the naris.

Figure 1. From Perner 2018, the original reconstruction of Wachtlerosaurus. Scale bar added.

Figure 1. From Perner 2018, the original reconstruction of Wachtlerosaurus. Scale bar added. Note the elongate ribs considered parts of the pelvis here. Pes of Euparkeria added by Perner. Manus appears to be from a coelophysoid theropod and flipped left to right.

The Dolomites are about 250 million years old
and are formed from coral reefs in the Tethys Sea, a perfect niche for a marine reptile like a thalattosaur.

Figure 1. Wachtlerosaurus in situ and reconstructed in lateral view.

Figure 2. Wachtlerosaurus in situ and reconstructed in lateral view.

A reconstruction of the skull helps
(Fig. 1) put the pieces of the broken skull back together again.

A few other new interpretations on the paper.

  1. Perner 2018 identifies two long, parallel dorsal ribs surrounding a jumble of ?vertebrae as parts of an oversized pelvis (Fig. 1).
  2. Perner employs the humerus in place of a scapula (Fig. 1).
  3. Probably this scattered bone specimen is incomplete, not nearly complete, as described.
Figure 3. Subset of the LRT nesting Wachtlerosaurus with Thalattosaurus.

Figure 3. Subset of the LRT nesting Wachtlerosaurus with Thalattosaurus.

References
Perner T 2018. A new interesting archosaur from the Ladinian (Middle Triassic) of the Dolomites (Northern Italy) Preliminary report. Pp 1–8 in Some new and exciting Triassic Archosauria from the Dolomites (Northern Italy). Perner T and Wachtler M eds. Dolomythos-Museum, Oregon Institute of Geological Research.

A new place to nest tiny Hadrocodium: with Morganucodon and Volaticotherium

I trusted published drawings
by Luo et al. 2001(Fig. 1) for tiny Hadrocodium and previously nested it with the tiny basalmost mammal, Megazostrodon (Fig. 2). That original drawing turned out to include a few errors. Here I trace and reconstruct Hadrocodium bones using DGS (Fig. 1). I also restored the dorsal skull shape to its invivo convex curve, which realigned the tooth rows with the jaw joint. I also recovered a down-turned premaxilla, relative to the maxilla, not an upturned one, as originally traced.

Figure 1. Hadrocodium skull in DGS tracing and reconstruction, plus the original drawing by Luo et al. 2001.

Figure 1. Hadrocodium skull in DGS tracing and reconstruction, plus the original drawing by Luo et al. 2001. The frontal/parietal hinge was pushed down during taphonomy. Here it is restored to its invivo position, which lines up the teeth with the jaw joint. Two molars nests it with other basal mammals with two molars.

Now
in the large reptile tree (LRT, 1258 taxa, not yet updated) Hadrocodium nests with other not-so-basal mammals, like UkhaatheriumMorganucodon and Volaticotherium, which we looked at yesterday. Like the latter two, Hadrocodium also has only two upper molars, a gracile zygomatic arch (jugal + squamosal) and other traits in common.

Hadrocodium wui (Luo, Crompton and Sun 2001), is known only from a very tiny skull. Due to its tiny size and lack of four to five molars (Fig. 2) Hadrocodium was originally considered a juvenile basal mammal, but later a tiny adult. Hadrocodium has a relatively larger brain size and more advanced ear structure than MegazostrodonHadrocodium nests with other taxa with two molars, Morganucodon and Volaticotherium.

Figure 1. Megazostrodon, an early mammal, along with Hadrocodium, a Jurassic tiny mammal.

Figure 2. Megazostrodon, an a Jurassic mammal, along with Hadrocodium, a Jurassic tiny mammal to scale and with Hadrocodium skull enlarged.

Revisiting basal mammals
is moving some of them around the LRT to nest with more similar sisters. There may be more such changes in the future as data improves. Things are a bit unsettled at present.

References
Luo Z-X, Crompton AW and Sun A-L 2001. New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics. Science 292 (5521): 1535–1540. Bibcode:2001Sci…292.1535L. doi:10.1126/science.1058476. PMID 11375489.

Where does the frigate bird nest?

For a blog focused on pterosaurs
it sure took a long time to take a look at the extant frigate bird (Figs. 1-3; genus: Fregata), a modern analog for many of the sea-faring clades of pterosaurs in terms of wing shape (long span, short chord) and gliding ability (see below).

FIgure 1. Fregata in flight.

FIgure 1. Fregata in flight. Note the narrow-chord wing membrane, as in all pterosaurs.

Between pelicans and cormorants 
In the large reptile tree (LRT, 1227 taxa) the frigate bird nests between the clade Pelecanus + Balaeniceps (the shoebill) and Phalacrocorax (the cormorant). The shoebill has the longest legs in the clade, so it is the most primitive member. Most studies, including the LRT and DNA analyses, associate frigate birds with pelicans, skuas and petrels, but some link frigate birds with a larger list including herons, ibises, spoonbills, hamerkops, penguins, loons, gannets, and cormorants. Why can’t DNA be more specific? That’s a wide gamut of taxa. The LRT is specific and fully resolved.

FIgure 2. Fregata skull with a closeup of the tiny jugal.

FIgure 2. Fregata skull with a closeup of the tiny jugal.

Interesting that frigate birds don’t like to get wet
while their sisters, cormorants dive for food, but then have to stand with wings dripping while drying out. Distinct from ducks, cormorant feathers don’t shed water with an oily coat.

Figure 4. Skeleton of Fregata, the frigate bird. Note the long bill, long neck and long antebrachium, perhaps the closest living analog to Cretaceous ornithocheirid pterosaurs.

Figure 3. Skeleton of Fregata, the frigate bird. Note the long bill, long neck and long antebrachium, perhaps the closest living analog to Cretaceous ornithocheirid and pteranodontid pterosaurs (Fig. 5). Consider this a shoebill stork and/or pelican with a slender bill and very short legs and you will be close to its phylogenetic grade.

Fregata magnificens (Lacépède, 1799; up to 56 cm long) inflates its throat sac with air, like a balloon, to display its bright red color (distinct from the pelican throat sac, which fills with water and prey). According to Wikipedia, “frigatebirds spend most of the day in flight hunting for food, and roost on trees or cliffs at night. The duration of parental care is among the longest of any bird species; frigatebirds are only able to breed every other year. Fossils date back to the Eocene, 50 mya.” 

Figure 2. Cearadactylus, Anhanguera and Pteranodon compared. The inset compares the humerus of Anhanguera and Pteranodon.

Figure 4. Cearadactylus, Anhanguera and Pteranodon compared. The inset compares the humerus of Anhanguera and Pteranodon. Compare proportions to the skeleton of Fregata. Look at those long wing tips, completely different from flightless pterosaurs, including large to giant azhdarchids. Most workers consider these taxa to be closely related, but the LRT does not confirm that.

Tested frigate birds
(Huey and Deutsch 2016) stayed aloft for two months without ever touching the ground riding cumulus thermals up to 6500 feet above sea level. 

Like ornithocheirid and pteranodontid pterosaurs
the torso is small and the wing has a narrow chord and a wide span in Fregata. This is also like modern man-made gliders, and unlike the proportions found in large to giant azhdarchids, which could not fly at all, contra traditional thinking, based on their relatively short distal wing finger phalanges (like those of small flightless pterosaurs).

References
Huey RB and Deutsch C 2016. How frigate birds soar around the doldrums. Science 353 (6294):26–27.
Lacépède BGÉ de 1799. Discours d’ouverture et de clôture du cours d’histoire naturelle : donné dans le Muséum national d’Histoire naturelle, l’an VII de la République, et tableaux méthodiques des mammifères et des oiseaux, Paris.

wiki/Frigatebird

Bergamodactylus (basal pterosaur) back ‘under the microscope’

This all started with Kellner 2015
who proposed 6 states of pterosaur ontogeny based on skeletal fusion of discrete elements. This hypothesis was tested in phylogenetic analysis and shown to be invalid. Pterosaurs don’t fuse bones during ontogeny. Fusion appears in phylogenic patterns. Oblivious to this fact, Dalla Vecchia 2018 dismissed Kellner’s hypothesis by writing, “Kellner’s six ontogenetic stages are an oversimplification mixing ontogenetic features of different taxa that probably had distinct growth patterns. Finding commonality across all pterosaurs is impossible, because there is much variation in pterosaur ontogeny and the available sample is highly restricted.” 

Neither Kellner nor Dalla Vecchia recognize
the lepidosaurian affinities of pterosaurs, and do not realize that as lepidosaurs pterosaurs mature differently than archosaurs. Some lepidosaurs continue growing after fusing elements (Maisano 2002). Others never fuse elements. Fusion of elements in pterosaurs is phylogenetic, not ontogenetic. Pterosaurs mature isometrically, not allometrically as proven by every full-term embryo and every known juvenile among a wide variety of pterosaur specimens. Plus, all of the small purported Solnhofen juveniles phylogenetically nest as key transitional taxa linking larger long-tail primitive pterosaurs to larger short-tail derived pterosaurs (Peters 2007). That’s how those clades survived the extinction events that doomed their fellow, larger, longer-tailed kin.

Kellner 2015 also
distinguished a small pterosaur MPUM 6009 from the holotype of Eudimorphodon and from Carniadactylus (MFSN 1797, Dalla Vecchia 2009; Fig. 1) and gave MPUM 6009 the name Bergamodactylus (Fig. 1) after Peters 2007 had done the same (without renaming MPUM 6009), in phylogenetic analysis. Neither Kellner nor Dalla Vecchia performed a phylogenetic analysis, but preferred to describe similar bones. That rarely works out well.

Figure 1. Bergamodactylus compared to Carniadactylus. These two nest apart from one another in the LRT.

Figure 1. Bergamodactylus (MPUM 6009) compared to Carniadactylus (MFSN 1797). These two nest apart from one another in the LRT. Contra Dalla Vecchia 2018, these two share relatively few traits in common. The feet, cervicals, sternal complex coracoids and legs are different.

Dalla Vecchia 2018 concludes, 
“The anatomical differences between MPUM 6009 and MFSN 1797 are too small to support the erection a new genus for MPUM 6009.” That is incorrect (Fig. 1). Several taxa nest between these two taxa in the large pterosaur tree (LPT, 232 taxa). Their feet alone (Fig. 1) were shown to be very different in Peters (2011).

Figure 1. Bergamodactylus compared to Cosesaurus. Hypothetical hatchling also shown.

Figure 2. Bergamodactylus compared to Cosesaurus. Hypothetical hatchling also shown.

From the Dalla Vecchia 2018 abstract
“Six stages (OS1-6) were identified by Kellner (2015) to establish the ontogeny of a given pterosaur fossil. These were used to support the erection of several new Triassic taxa including Bergamodactylus wildi, which is based on a single specimen (MPUM 6009) from the Norian of Lombardy, Italy. However, those ontogenetic stages are not valid because different pterosaur taxa had different tempos of skeletal development. Purported diagnostic characters of Bergamodactylus wildi are not autapomorphic or were incorrectly identified. Although minor differences do exist between MPUM 6009 and the holotype of Carniadactylus rosenfeldi, these do not warrant generic differentiation. Thus, MPUM 6009 is here retained within the taxon Carniadactylus rosenfeldi as proposed by Dalla Vecchia (2009a).” \

Dalla Vecchia is basing his opinion on comparing a few cherry-picked traits, possibly convergent, rather than running both taxa and a long list of other pterosaurs through phylogenetic analysis, to see where unbiased software nests both taxa among the others.

Plus, as mentioned above, both authors are working from an antiquated set or rules that no longer apply now that pterosaurs have been tested and validated as lepidosaurs.

Figure 2. Bergamodactylus skull colorized with DGS and reconstructed.

Figure 3. Bergamodactylus skull colorized with DGS and reconstructed. Palatal and occipital bones shown here were missed by Dalla Vecchia 2018 and prior workers who did not use DGS.

Phylogenetic analysis
employing a large gamut of taxa, like the large reptile tree (LRT, 1215 taxa), invalidates traditional arguments that pterosaurs arose without obvious precedent among the archosauriforms, which most pterosaur workers, including both Kellner and Dalla Vecchia, still cling to, despite no evidence of support. Pterosaurs arose from fenestrasaur tritosaur lepidosaurs (Fig. 7).

Figure 4. The skull of Bergamodactylus traced by Kellner 2015, Dalla Vecchia 2018 and by me using DGS.

Figure 4. The skull of Bergamodactylus traced by Kellner 2015, Dalla Vecchia 2018 and by me using DGS. See figure 2 for a reconstruction of the DGS tracing.  Prior authors missed all the palatal and occipital bones along with several others.

The metacarpus of Bergamodactylus
has a few disarticulated elements. When replaced to their in vivo positions the axial rotation of metacarpal 4 (convergent with the axial rotation of pedal digit 1 in birds) enables the wing finger to fold in the plane of the hand, not against the palmar surface. Manual digit 5, a vestige, goes along for the ride, rotating the dorsal surface of the hand (Fig. 5).

Figure 5. Metacarpus of Bergamodactylus (MPUM 6009) in situ and reconstructed.

Figure 5. Metacarpus of Bergamodactylus (MPUM 6009) in situ and reconstructed. Apparently the pteroid splintered apart, overlooked by those with direct access to the specimen. The distal carpals are not co-ossified, as they are in later pterosaurs. The laterally longer fingers, up to digit 4, is a tritosaur trait. Note ungual 1 lies on top of the posterior face of metacarpal 4. That was overlooked by those who had direct access to the specimen, which supports the utility of DGS.

 

Bergamodactylus, as the most basal pterosaur,
is itself a transitional taxon bridging non-volant fenestrasaurs with all other pterosaurs. And the wing (Fig. 6) was about the last thing to evolve.

Figure 6. Click to enlarge. The origin of the pterosaur wing and the migration of the pteroid and preaxial carpal. A. Sphenodon. B. Huehuecuetzpalli. C. Cosesaurus. D. Sharovipteryx. E. Longisquama. F-H. The Milan specimen MPUM 6009, a basal pterosaur.

Bergamodactylus to scale
with Cosesaurus and Longisquama (Fig. 7), demonstrate the variety within the Fenestrasauria. Pterosaurs arose more or less directly from a sister to Cosesaurus (based on overall proportions), but note that both Sharovipteryx and Longisquama have more pterosaurian traits than Cosesaurus does. This pattern is convergent with that of birds, of which several clades of Solnhofen bird descendants arose of similar yet distinct structure.

Figure 8. Taxa at the genesis of pterosaurs: Cosesaurus, Longisquama and Bergamodactylus.

Figure 7. Taxa at the genesis of pterosaurs: Cosesaurus, Longisquama and Bergamodactylus.

See rollover images
of Bergamodactylus in situ here. You’ll see how DGS is able to pull out post-cranial details overlooked by others in the chaos and confusion of layers of bones and impressions in MPUM 6009. Cranial details are best seen in figure 3 above, which is based on higher resolution images.

References
Dalla Vecchia FM 2009. Anatomy and systematics of the pterosaur Carniadactylus gen. n. rosenfeldi (Dalla Vecchia, 1995). Riv. It. Paleontol. Strat., 115: 159-186.
Dalla Vecchia FM 2018. Comments on Triassic pterosaurs with a commentary on the “ontogenetic stages” of Kellner (2015) and the validity of Bergamodactylus wildi.  Rivista Italiana di Paleontologia e Stratigrafia 124(2): 317-341. DOI: https://doi.org/10.13130/2039-4942/10099 https://riviste.unimi.it/index.php/RIPS/article/view/10099
Kellner AWA. 2015. Comments on Triassic pterosaurs with discussion about ontogeny and description of new taxa. Anais Acad. Brasil. Ciênc., 87(2): 669-689.
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrate Paleontology 22:268-275.
Peters D. 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification. Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605

4 nostrils in Chamaeleo?

The skull of the smooth chameleon,
Chamaeleo laevigatus (Figs. 1, 2), has two extra holes in the anterodorsal plane of its rostrum (Fig. 1). Despite appearances, the holes visible in top view are not nostrils.

Figure 1. The chameleon Trioceros jacksonii colored using DGS. The sutures are difficult to see in the original skull, much easier in the colorized tracing.

Figure 1. The chameleons Chamaeleo and Trioceros. Note the lateral nostrils on both taxa. Chamaeleo has two more openings in dorsal view.  Not sure if Trioceros was the same. Note the giant pterygoids on Chamaeleo. The prefrontal and postfrontal are in contact. The premaxilla is tiny in ventral view.

The Chamaeleo rostrum
is angled at about 50º from the jawline. Given just the skull, you might think those openings in dorsal view are nostrils. With skin and scales on (Fig. 2), the nostrils are located on the lateral plane, as in other chameleons, like Trioceros (Fig. 1), surrounded by traditional circumnarial bones.

Figure 2. Chamaeleo laevigatus invivo. Red arrow points to external naris.

Figure 2. Chamaeleo laevigatus invivo. Red arrow points to external naris.

Diaz and Trainer 2015 published
some nice images of chameleon hands and feet, colorized here (Fig. 3) for additional clarity. The metacarpals and metatarsals are the bones that radiate. The phalanges are all vertical here.

Figure 3. The manus and pes skeleton of a chameleon from Diaz et al. 2016 with colors added and the second from left image relabels the fingers, correcting a typo.

Figure 3. The manus and pes skeleton of a chameleon from Diaz et al. 2015 with colors added and the second from left image relabels the fingers, correcting a typo. Manual 1 has only two phalanges. The metacarpals and metatarsals open horizontally in these images. Note the ankle elements are not co-ossified.

References
Diaz RE Jr. and Trainor PA 2015. Hand/foot splitting and the ‘re-evolution’ of mesopodial skeletal elements during the evolution and radiation of chameleons. BMC Evolutionary Biology201513:184.

wiki/Smooth_chameleon
digimorph.org/Chamaeleo_laevigatus/
Chamaeleo laevigatus GRAY, 1863″. The Reptile Database

Monodon: THE weirdest skull of all mammals

Today two blogposts are published
because they relate strongly to one another. Here is the post on torsioned tenrec/odontocete skulls.

Figure 1. Distinct from most narwhals, this skull also has right tusk emerging from the canine position. And yes, that's the maxilla covering most of the skull, even above the orbit! I added an eyeball here to help locate the orbit. The mesethmoid is the red bone that divides the naris (blow hole).

Figure 1. Distinct from most narwhals, this skull also has right tusk emerging from the canine position. And yes, that’s the maxilla covering most of the skull, even above the orbit! I added an eyeball here to help locate the orbit. The mesethmoid is the red bone that divides the naris (blow hole).

The narwhal (genus Monodon, Fig. 1)
is famous for having one giant spiral tooth sticking out ahead of its skull. Monodon also has one of the most bizarre skulls of all mammals and departs from that of all tetrapods, partly due to the root of the tooth and partly due to the migration of the nares to the back of the skull. Except for its tips, the jugal is missing. The maxilla, lacks teeth (if you don’t count the tusk) and rather than extending below the orbit, extends over the forehead, following the naris on its migration to the back of the skull. The bulbous portion of the skull, the cranium is made of parietals in most mammals, but the parietals are greatly reduced, nearly absent in Monodon.

Figure 2. The beluga, Delphinapterus, is closely related to, though less derived than the narwhal, Monodon. More teeth of a regular shape were present in the jaws. Those two yellow arrows indicate a misalignment of the centerline of the top of the occiput vs. the bottom. Compare to figure 3.

Figure 2. The beluga, Delphinapterus, is closely related to, though less derived than the narwhal, Monodon. More teeth of a regular shape were present in the jaws. Those two yellow arrows indicate a misalignment of the centerline of the top of the occiput vs. the bottom. Compare to figure 3. The mesethmoid is the red bone in the blow hole. This skull is also bent left, as in the narwhal.

The sister taxon of the narwhal
is the beluga (genus: Delphinapterus). It helps one understand the narwhal a bit better. At least the beluga has a few traditional teeth. These two taxa nest together in the large reptile tree (LRT, 1087 taxa, Fig. 4).

Figure 3. Chonecetus has a more primitive skull with nares closer to the snout tip and no maxilla above the orbit.

Figure 3. Chonecetus has a more primitive skull with nares closer to the snout tip and no maxilla above the orbit. Not a transitional taxon to baleen whales, which have another separate origin. This drawing lacks any indication of torsion, perhaps because the back half was separated from the front half and the artist ‘repaired’ the twist.

Less derived and more primitive
is Chonecetus (Fig. 3), which has nares closer to the snout tip, and more teeth, and more cranium. This taxon and its sister, Aetiocetus, have been traditionally considered transitional from toothed whales to baleen whales, like Balaenoptera, but baleen whales have an entirely separate ancestry derived from desmostylians, like Desmostylus.

Figure 5. Subset of the LRT focusing on the tenrec/odontocete clade with several whales added.

Figure 4. Subset of the LRT focusing on the tenrec/odontocete clade with several whales added.

A recent paper on Monodon tusks (Nweeia et al. 2012)
found “the narwhal tusks are the expression of canine teeth and that vestigial teeth have no apparent functional characteristics and are following a pattern consistent with evolutionary obsolescence.” (See Figs. 5, 6).

Figure 4. Image from Nweeia et al. 2012 showing the unerupted right tusk and the root of the left tusk in the male narwhal along with two unerupted tusks in the female.

Figure 5. Image from Nweeia et al. 2012 showing the unerupted right tusk and the root of the left tusk in the male narwhal along with two unerupted tusks in the female. Note the angle of the posterior skull relative to the anterior midline.

In dorsal or ventral view
it is clear that the the tusk (left) side of the skull is longer than the right side due to angling the posterior skull relative to the rostrum.

Figure 6. CT scans of a female narwhal (Monodon) showing soft tissues and unerupted teeth. Note the angle of the posterior skull relative to the anterior.

Figure 6. CT scans of a female narwhal (Monodon) showing soft tissues and unerupted teeth. Note the angle of the posterior skull relative to the anterior. The left side, the tusk side, is shorter than the right side in figure 5, so the label ‘ventral’ is an error here. This is a dorsal view of the female skull in figure 5. Always test scale bars and labels.

I wonder about the bending of the skull
toward the left in these two whales. Could asymmetry have anything to do with stereo auditory senses? Asymmetry is also found in owl skulls, another taxon that depends strongly on acute hearing for locating prey.

Figure 7. Fetal narwhal skull, here colorized from Nweenia et al. 2012. The jugal disappears in adults.

Figure 7. Fetal narwhal skull, here colorized from Nweenia et al. 2012. The jugal disappears in adults. The asymmetry is already apparent here.

Figure 8. Common bottle nose dolphin skull (genus: Tursiops) also displays a bit of asymmetry in dorsal view.

Figure 8. Common bottle nose dolphin skull (genus: Tursiops) also displays a bit of asymmetry in dorsal view. Note the yellow arrows on the parietal showing the wee bit of torsion here. 

Update:
With 1187 taxa and 231 traits full resolution was recovered in the LRT after running PAUP FOR 16 minutes and 15 seconds. The single best tree has 16,329 steps.

References
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.’
Nweeia MT et al. (9 co-authors) 2012. Vestigial tooth anatomy and tusk nomenclature for Monodon monoceros. The Anatomical Record 295:1006–1016.
Pallas PS 1766. Miscellanea Zoologica.

wiki/Narwhal
wiki/Beluga_whale