Is Thecocoelurus an ornithomimosaur?

Allain et al. (2014) recently claimed Thecocoelurus, a small partial theropod cervical vertebra from Early Cretaceous England, is "morphologically identical" to an undescribed supposed ornithomimosaur from the Early Cretaceous of France (from here on called the Angeac taxon).  After reading Andrea Cau's post on the topic, I decided to do some in depth comparison.

Besides the Angeac taxon, Thecocoelurus has been compared to several taxa with surprisingly similar cervical vertebrae- caenagnathids (Naish et al., 2001; Naish and Martill, 2002); Falcarius (Kirkland et al., 2004; Zanno, 2010a); and noasaurids (Naish, 2011).  These all have elongate amphicoelous cervicals with low neural spines, anterior peduncular fossae, two pairs of pleurocoels and a transversely concave ventral surface defined by lateral ridges confluent with the parapophyses.  Which is most similar to Thecocoelurus?

Naish et al. (2001) and Naish and Martill (2002) both argue it is closer to oviraptorosaurs than to therizinosauroids based on the rounded pleurocoel and thin neural spine, but this is also the case in basal therizinosaurs (Falcarius, Jianchangosaurus) (as noted by Zanno, 2010a) and noasaurids. Naish (2011) on the other hand, felt "the idea that large caenagnathids were present in Western Europe during the Barremian is difficult to reconcile with what we know of oviraptorosaur biogeography and distribution", thus favored an abelisauroid identity (I suppose based on Genusaurus).  I'd argue basal oviraptorosaurs could and do have similar morphologies (e.g. the Early Cretaceous Similicaudipteryx) and that small theropod diversity in Cretaceous Europe is very poorly known. 

Allain et al. make two new comparative arguments for Thecocoelurus being closest to the Angeac taxon.
1. They distinguished both from Noasaurus based on their concave anterior central surface, but this is true in Masiakasaurus as well. It is also true of all coelurosaurs being compared.
2. They also paired Thecocoelurus with the Angeac taxon based on pneumatic foramina above the prezygapophyses which invade the neural arch. Yet these are present in cervicals 6-10 of Masiakasaurus (Allain et al. state they are "in a modified form" but don't elaborate), at least cervical 9 of Heyuannia, and in Conchoraptor and "Ingenia" (Lu, 2004; contra Allain et al.'s claim they are unknown in oviraptorosaurs). They are also only present on the left side of Thecocoelurus (pf? in figure below, lower left), further posterior than in at least Masiakasaurus and the Angeac taxon, perhaps suggesting breakage of a naturally hollow area or pneumatic asymmetry. The only preserved posterior cervical of Falcarius doesn't preserve this area, nor does the illustrated and best preserved cervical of Chirostenotes.

So let's compare!  Contra Naish and Martill, the specimen resembles posterior cervicals more than anterior ones, and indeed the supposedly identical Angeac vertebra matches the seventh or eighth of Harpymimus based on elongation, central articular surface orientation, prezygapophyseal length and orientation, etc. (contra Allain et al.).

Anterior half of posterior cervical vertebrae in (descending) right lateral, anterior, ventral and dorsal views.  From left to right- Thecocoelurus holotype (after Naish and Martill, 2002); Angeac taxon ANG 10/175 (after Allain et al., 2014); Falcarius UMNH VP 14657 (after Zanno, 2010b); Masiakasaurus FMNH PR 2481 (after Carrano et al., 2011); Chirostenotes or Epichirostenotes ROM 43250 (after Sues, 1997); Similicaudipteryx holotype (after He et al., 2008).

The elongate parapophyses resemble Falcarius, Chirostenotes and Similicaudipteryx more than Masiakasaurus or the Angeac taxon.
The anterior pleurocoels are placed in an obvious fossa, like Falcarius and the Angeac taxon, but unlike Masiakasaurus, Chirostenotes or Similicaudipteryx.
The infradiapophyseal fossa is developed as an elongate groove, as in Falcarius but unlike Masiakasaurus, the Angeac taxon, Chirostenotes or Similicaudipteryx.
The centrum is taller than wide (midline height / width minus parapophyses 133%) as in Falcarius (118%), but unlike the Angeac taxon (95%) and especially Chirostenotes (74%) and Masiakasaurus (64%).
The anterior peduncular fossae are well defined as in Chirostenotes and at least anterior Falcarius cervicals, but unlike Masiakasaurus or the Angeac taxon. They are however also well defined in anterior Masiakasaurus cervicals, so the condition in Falcarius isn't necessarily better than that genus or the Angeac taxon.
They are also placed far below the diagonal prezygapophyseal surface as in at least anterior Falcarius cervicals, but unlike the Angeac taxon, Chirostenotes or Masiakasaurus. The same could be said re: anterior Masiakasaurus cervicals.
The prespinal fossa is broad like Chirostenotes and Masiakasaurus but unlike the Angeac taxon.
It has anteroposteriorly broad exposure dorsally as in the Angeac taxon and at least anterior Falcarius cervicals, but not Masiakasaurus (including anterior cervicals of the latter).

Overall, Thecocoelurus is most similar to Falcarius and least similar to Masiakasaurus.  There are four good characters shared with Falcarius to the exclusion of the Angeac taxon, and three characters that are more similar to Chirostenotes than to the Angeac taxon, but two characters that are more similar to the Angeac taxon than to Chirostenotes.

Thecocoelurus holotype completed with the posterior of Falcarius (modified from Naish and Martill, 2002 and Zanno, 2010).  This results in a centrum length of 68 mm for Thecocoelurus, compared to Naish's (2011) estimate of 70-90 mm.

Falcarius does differ from Thecocoelurus in having a ventral median ridge on its centra, but this is an autapomorphy not seen in other therizinosaurs.  Besides this, no characters differ between the specimen except exact size and shape of pneumatic features, which themselves vary between right and left sides of Thecocoelurus.  Both are Barremian, and Thecocoelurus is 58% the size of the Falcarius individual that preserved the posterior cervical (though a growth series is known, where that individual falls is unreported).  Whether Thecocoelurus and Falcarius share derived characters to the exclusion of other therizinosaurs would require more study, but at the moment is seems most parsimonious to consider Thecocoelurus a basal therizinosaur and not closely related to the Angeac taxon.

References- Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from Western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.

Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. In Martill and Naish (Eds). Dinosaurs of the Isle of Wight. The Palaeontological Association. 242-309.

Naish and Martill, 2002. A reappraisal of Thecocoelurus daviesi (Dinosauria: Theropoda) from the Early Cretaceous of the Isle of Wight. Proceedings of the Geologists’ Association. 113, 23-30.

Kirkland, Zanno, DeBlieux, Smith and Sampson, 2004. A new, basal-most therizinosauroid (Theropoda: Maniraptora) from Utah demonstrates a Pan-Laurasian distribution for Early Cretaceous therizinosauroids. Journal of Vertebrate Paleontology. 24(3), 25-26.

Lu, 2004. Oviraptorid dinosaurs from Southern China. PhD Thesis. Southern Methodist University. 249 pp.

He, Wang and Zhou, 2008. A new genus and species of caudipterid dinosaur from the Lower Cretaceous Jiufotang Formation of Western Liaoning, China. Vertebrata PalAsiatica. 46(3), 178-189.

Zanno, 2010a. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic Palaeontology. 8(4), 503-543.

Zanno, 2010b. Osteology of Falcarius utahensis: Characterizing the anatomy of basal therizinosaurs. Zoological Journal of the Linnaean Society. 158, 196-230.

Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001, and implications for the morphology of the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology. 95, 53 pp.

Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The Palaeontological Association. 526-559.

Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria, Theropoda): An undetected record. Geologica Acta. 12(2), in press. http://www.geologica-acta.com/pdf/vol1202a05.pdf

Revisiting a BAND cladogram

Twelve years ago, I replied to a post Nick Gardner wrote to the DML, critiquing and updating the phylogenetic analysis of Hou et al. (1996).  This was perhaps the first analysis that I corrected and reran, which is now the concept behind an entire section of my website.  Well, now I've officially reexamined Hou et al.'s data for an entry in the Database.

To jog everyone's memory, Hou et al.'s paper provided details on Confuciusornis, Liaoningornis and Chaoyangia and was interesting in that it included a cladogram with matrix for birds, not before (or since?) done by the Birds Are Not Dinosaurs (BAND) crowd.  This cladogram supported the BAND concept of Sauriurae including Confuciusornis and enantiornithines, and supposedly used Petrolacosaurus as an outgroup.  Hou et al. claim "The use of a coelurosaurian dinosaur like Velociraptor as the outgroup would make no difference for character states."  Really?  The figured cladogram also oddly differed from the matrix in  adding 'Modern Birds' as sister to Ichthyornis and replacing Cathayornis with Enantiornithes.  The figure has a similar but different character list than the matrix (note my old post dealt with the figure's character list, so the numbers don't match up with my new critique).  The authors' phylogeny was-

|--outgroup (Petrolacosaurus)
`--Aves
   |--Sauriurae
   |  |--Archaeopteryx
   |  `--+--Confuciusornis
   |     `--Cathayornis
   `--Ornithurae
      |--Liaoningornis
      `--+--Chaoyangia
         `--+--Hesperornis
            `--Ichthyornis

Correcting and re-running the matrix

Go here to read commentary on all of the characters and codings.  This analysis is hindered by its small size, several characters based on fictional morphologies, others that are the opposite of other characters, and some which are based on developmental assumptions that cannot be coded in adult specimens. This leaves only seventeen characters that are valid and phylogenetically informative. The anatomy of Archaeopteryx has been greatly misunderstood by BANDits for decades, which spread somewhat to Confuciusornis. Hou et al. also ascribed unsupported referred specimens to Cathayornis and Chaoyangia, negating many of the codings for those taxa. Petrolacosaurus was coded 0 for everything, including characters it actually has state 1 for, those which are unpreserved, and even those based on structures it lacks. Ichthyornis meanwhile was coded as if it were a complete ornithurine, despite not preserving several elements. These examples show Hou et al. coded taxa as idealized hypothetical examples instead of actual specimens. In total, 43% (110/256) of characters were miscoded, though this is somewhat exaggerated by the several deleted characters. Once corrected, the consensus is-

|--Petrolacosaurus
`--Eosuchia to Ornithes
   |--Archaeopteryx
   `--Ornithurae
      |--Enantiornithes
      |  |--Cathayornis
      |  `--Liaoningornis
      `--Pygostylia
         |--Confuciusornis
         `--+--Chaoyangia
            `--Ornithuromorpha
               |--Hesperornis
               `--Ichthyornis

This differs from the original tree in having a paraphyletic 'Sauriurae', though differs from the modern consensus in placing Confuciusornis closer to Aves than enantiornithines. This is no doubt due to the low number of characters, and as noted on my site, changed with only 2 extra steps. Liaoningornis moved to Enantiornithes, matching O'Connor's (2012) redescription.  Hou et al.'s original flawed matrix finds Pygostylia and Ornithothoraces instead of Sauriurae with only 3 extra steps, so doesn't even support their own hypothesis strongly.  Indeed, my conclusion is that the matrix is too small to strongly reject any proposed hypothesis.

What about Petrolacosaurus?

It's very odd that Hou et al. would choose Petrolacosaurus as their avian outgroup.   I mean, even BANDits think birds are archosaurs, so why not use Euparkeria or Ornithosuchus?  As noted above, they state "The use of a coelurosaurian dinosaur like Velociraptor as the outgroup would make no difference for character states" compared to their supposed use of Petrolacosaurus. Well let's check that.

If Velociraptor is added, there are in fact nine characters that are coded differently. Velociraptor is known to have feathers (based on ulnar quill knobs) whereas Petrolacosaurus' integument is unpreserved though originally miscoded as known.  Petrolacosaurus is inapplicable for hypocleidium length, anterior sternal groove presence and sternal length (all originally miscoded), due to lacking a furcula or ossified sternum, unlike Velociraptor. Velociraptor has uncinate processes, unlike Petrolacosaurus or even Archaeopteryx. Velociraptor has an ectocondylar tuber on its femur like Archaeopteryx, though Hou et al. view the latter's structure as a non-homologous structure. Petrolacosaurus' pedal unguals are not enlarged (again miscoded by Hou et al.), but Velociraptor has pedal ungual II enlarged so is polymorphic. Most amusingly, Petrolacosaurus actually has a dorsolaterally grooved clavicle and proximodorsal ischial process like some basal birds (but unlike Velociraptor), though Hou et al. miscoded these in assuming it was a generic diapsid.  That's right.  The BANDits had data to support their hypothesis, but didn't realize it due to their own laziness and evolutionary assumption that these were derived sauriurine characters instead of symplesiomorphies.

Bird-like characters of Petrolacosaurus assumed to be absent by Hou et al. (1996).  Left- left clavicle in lateral view showing posterolateral groove in pink. Right- left pelvis in lateral view showing proximodorsal ischial process in pink. Both modified after Reisz (1981).

Incidentally, if Velociraptor replaces Petrolacosaurus, the tree is the same except that confuciusornithids, enantiornithines and ornithuromorphs are in a trichotomy, allowing the current consensus.

References- Reisz, 1981. A diapsid reptile from the Pennsylvanian of Kansas. University of Kansas Museum of Natural History. Special Publication 7, 74 pp.

Hou, Martin, Zhou and Feduccia, 1996. Early adaptive radiation of birds: Evidence from fossils from northeastern China. Science. 274, 1164-1167.