A quick look at the original Tapejara skull

Short one today
told mostly in pictures.

Topic: Simplification. 
Get rid of the extraneous data to better see and understand the basics.

Figure 1. Just take out everything that isn't this side of Tapejara. Rearrange the parts for best fit. Make a guess for the missing parts and this is what you get.

Figure 1. Just take out everything that isn’t this side of Tapejara. Rearrange the parts for best fit. Make a guess for the missing parts and this is what you get.

In this case
the skull of the original Tapejara fossil (Fig. 1) is trimmed back to just the basics and slightly shifted to fit. Missing tips are added based on phylogenetic bracketing.

Figure 4. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

Figure 2. Click to enlarge. Tiny Tapejaridae arise from dsungaripterids and germanodactylids, then grow larger phylogenetically.

Tapejara wellnhoferi
(Kellner 1989; 108 mya, Early Cretaceous) was immediately recognized as something quite different when first discovered. Compared to other specimens, this one appears to have no sharp premaxilla, likely due to taphonomic loss.


References
Eck K, Elgin RA and Frey E 2011. On the osteology of Tapejara wellnhoferi KELLNER 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil. Swiss Journal of Palaeontology, doi:10.1007/s13358-011-0024-5.
Kellner AWA 1989. A new edentate pterosaur of the Lower Cretaceous from the Araripe Basin, northeast Brazil. Anais da Academia Brasileira de Ciências 61, 439-446.

wiki/Tapejara

 

Shedding new light (literally!) on Jianianhualong: Li et al. 2020

Li et al. 2020 used various frequencies of light
and spectroscope technology on the holotype bones and feathers of Jianianhualong (Figs. 1, 2; Early Cretaceous, Xu et al. 2020, DLXH 1218) to identify specific elements in the matrix and specimen.

From the abstract:
“Here, we carried out a large-area micro-X-Ray fluorescence (micro-XRF) analysis on the holotypic specimen of Jianianhualong tengi via a Brucker M6 Jetstream mobile XRF scanner.”

Figure 2. Jianianhualong, Serikornis and Jurapteryx to scale.

Figure 1a. Jianianhualong, Serikornis and Jurapteryx to scale.

Figure 1. Jianianhualong tengi in situ. This is the largest among the early birds, a fact overlooked by the Xu et al. 2017. Think of Jianianhualong as a giant Archaeopteryx!

Figure 1b. Jianianhualong tengi in situ. This is the largest among the early birds, a fact overlooked by the Xu et al. 2017. Think of Jianianhualong as a giant Archaeopteryx!

From the abstract:
“Jianianhualong tengi is a key taxon for understanding the evolution of pennaceous feathers as well as troodontid theropods, and it is known by only the holotype, which was recovered from the Lower Cretaceous Yixian Formation of western Liaoning, China.” 

What they didn’t do is to rerun their phylogenetic analysis with more taxa (Fig. 2).

What they didn’t do is to create a reconstruction, perhaps using DGS to precisely trace and segregate the bones to rebuild the skeleton (Figs. 1, 3, 4).

Figure 2. Subset of the LRT focusing on Pennaraptora 2014 = Tyrannoraptora 1999. Here Khaan and Velociraptor substitute for Oviraptor and Deinonychus.

Figure x. Subset of the LRT focusing on birds and their ancestors. Jianianhualong nests within Aves (five taxa from the bottom).

By contrast,
in the large reptile tree (LRT, 1730+ taxa) Jianianhualong nests within Aves (five taxa from the bottom of Fig. 2) even though it was clearly not volant due to its much larger size and smaller forelimbs. Close relatives include Archaeopteryx (= Jurapteryx) recurva (= Eichstätt specimen, Fig. 3) and the privately held #11 specimen of Archaeopteryx.

The authors think Jianianhualong is a troodontid.
According to Wikipedia“A number of characteristics allow Jianianhualong to be identified as a member of the Troodontidae. These include:

  1. the long forward-projecting branch and flange of the lacrimal bone; [✓]
  2. the foramina on the nasal bone; [?]
  3. the smooth transition between the eye socket and the backward-projecting branch of the frontal bone; [✓]
  4. the ridge on the forward-projecting branch of the jugal bone; [✓]
  5. the triangular dentary bearing a widening groove; [✓]
  6. the robust forward-projecting branch of the surangular bone; [✓]
  7. the relatively large number of unevenly-distributed teeth; [✓]
  8. the flattened chevrons with blunt forward projections and bifurcated backward projections; [✓]
  9. and the broad and flat “pubic apron” formed by the pubic bones.” [?]
Figure 3. The Eichstätt specimen, Jurapteryx recurva, nests with the living ostrich, Struthio, presently in the LRT.

Figure 2. The Eichstätt specimen, Jurapteryx recurva, nests with the living ostrich, Struthio, presently in the LRT.

Professor Larry Martin would be so proud!
Why? Because the Wikipedia author (above) is using a list of traits to support an hypothesis of interrelationships rather than using a cladogram to support that hypothesis.  Checkmarks [✓] indicate traits Jurapteryx shares. Question marks [?] indicate traits not shown in Jianianhualong or Jurapteryx. Or did I miss something?

The problem is,
various authors add taxa to the Troodontidae that don’t belong there in the LRT, as we learned earlier here. The LRT; subset Fig. x) recovers Jiaianhualong as the largest known member of the Sapeornis/Jurapteryx clade of birds. Several flightless birds are in this clade. These could be confused with troodontids for that reason. In the LRT the clade Troodontidae include Sinornithoides + Sauronithoides their LCA and all derived taxa. None of these are direct bird ancestors.

Getting back to chemistry
“The bone in Jianianhualong is, as expected rich in calcium and phosphorus, corresponding mineralogically to apatite. The regions where feather remains can be observed show an enrichment and correlation pattern of several elements including manganese, titanium, nickel and copper.”

FIgure 2. GIF animation of the skull of Jianianhualong showing original tracing in line art and colorized bones (DGS) used to create a reconstruction (Fig. 3).

FIgure 3. GIF animation of the skull of Jianianhualong showing original tracing in line art and colorized bones (DGS) used to create a reconstruction (Fig. 3).

Jianianhualong is a troodontid-like bird,
not a bird-like troodontid. Note the odd scapula shape, like that in Sapeornis. Note the retrovered pedal digit 1, showing this taxon was derived from perching birds. The tall naris and long tibia are autapomorphies.

Xu et al. 2014 made a headline out of
the asymmetric feathers found with Jianianhualong. In the present context, Jianianhualong is derived from volant ancestors. So asymmetry is expected, not exceptional. This is the earliest known large flightless bird, not an example of the invalid hypothesis of ‘mosaic’ evolution.

Figure 3. Reconstruction of the skull of Jianianhualong based on DGS tracings in figure 2.

Figure 4. Reconstruction of the skull of Jianianhualong based on DGS tracings in figure 2.

Liaoningventor curriei (Shen et al. 2017; DNHM D3012; Early Cretaceous) was also originally described as a non-avian troodontid, but nests with Jianianhualong as a flightless bird.


References
Li J, et al. (8 co-authors 2020. Micro-XRF study of the troodontid dinosaur Jianianhualong tengi reveals new biological and taphonomical signals. bioRxiv 2020.09.07.285833 (preprint) PDF doi: https://doi.org/10.1101/2020.09.07.285833
https://www.biorxiv.org/content/10.1101/2020.09.07.285833v1
Shen C-Z, Zhao B, Gao C-L, Lü J-C and Kundrat 2017. A New Troodontid Dinosaur (Liaoningvenator curriei gen. et sp. nov.) from the Early Cretaceous Yixian Formation in Western Liaoning Province. Acta Geoscientica Sinica 38(3):359-371.
Xu X, Currie P, Pittman M, Xing L, Meng QW-J, Lü J-C, Hu D and Yu C-Y 2017. Mosaic evolution in an asymmetrically feathered troodontid dinosaur with transitional features. Nature Communications DOI: 10.1038/ncomms14972.

wiki/Sapeornis
wiki/Jianianhualong
wiki/Liaoningvenator

Variations on a Cutleria Restoration

Crushed fossils missing some of the middle/connecting bones give paleontologists several options for restoration. Case in point: Cutleria (Figs. 1, 2) a very basal therapsid. We looked at this specimen/taxon earlier here and here.

Figure 1. Cutleria restoration version 1, straight maxilla.

Figure 1. Cutleria restoration version 1, straight maxilla. Click to enlarge.

The first restoration aligns the ventral maxilla in a straight line. Note the very deep reflected lamina on the angular, sharply angled from the dentary.

Figure 2. Cutleria restoration version 2, convex maxilla.

Figure 2. Cutleria restoration version 2, convex maxilla. Click to enlarge.

The second restoration (Fig. 2) is the more conservative (more like the in situ fossil) producing a deeper, ventrally convex maxilla. Note the reflected lamina of the angular is not so deep here. In both cases the mandible needs to mate to the maxilla, so it changes in each case, enabled by the mid-length break.

Which one is more correct?

Factors to consider:

  1. Ancestral taxa (like Haptodus and Ophiacodon) have a shallow reflected lamina.
  2. Descendant taxa (like Biarmosuchus) have a deeper reflected lamina; but then
  3. Other descendant taxa (like Stenocybus) have a shallow reflected lamina.
  4. Descendant taxa (like Haptodus and Ophiacodon) have a convex maxilla.
  5. Descendant taxa (like Biarmosuchus have a less convex to straight maxilla.
  6. Other descendant taxa (like Stenocybus, dromasaurs, dicynodonts and other therapsids) have a convex maxilla.
  7. Version 2 requires less bone movement.
  8. Version 1 requires less ‘putty’ to fill in holes left by missing bones. The puzzle pieces fit tighter.
  9. The strong posterior lean of the lateral temporal fenestra is not present in either ancestral or descendant taxa and it probably reflects the angle of the mandible adductor.

Your opinion or insights would be appreciated.