Peters' theropod craziness

So David Peters, whose analyses I have thoroughly trashed on this blog before, has branched out into theropods.  In my prior foray into his basal dinosaurian phylogeny, I was surprised by how poorly coded (~30% miscoded) taxa were for his tiny and badly formed analysis.  Now that he's trying to interpret MY turf, it's just sad.  Or laughable.  Or laughably sad, take your pick.  Even in the best of circumstances, taking 228 characters designed for amniote phylogeny and coding theropods for them is likely to give you a terrible result.  But add Peters' DGS to the mix, where he takes photos and discovers missing bones in a slab or traces his own outlines, and you have hilarious fiction.  Peters might know pterosaurs, but when he tries tracing theropods, it's obvious he does NOT know theropods.  He mixes bone outlines, identifies random sediment texture or feathers as bones, and based on his results just has no idea how theropods look.  I commented on this on his Archaeornithura skull post last week (see last figure below), but Peters deleted the comment and changed his reconstruction to be wrong in different ways (new ways include that the pubis is the ischium, the ischium has a huge obturator process, the manus has a complete digit, as does the fifth pedal digit(!)).

Reconstructions of Sinosauropteryx prima by me (top), and Peters (bottom), scaled to same femoral length.

As an example, here's Sinosauropteryx as reconstructed by him vs. by me.  Now, Peters is no doubt the more experienced artist, and my reconstruction isn't perfect.  I made it back in 2005, and if I were to redraw it today, I'd give the tibia and fibula less generic shapes (e.g. larger cnemial crest, flatter distal tibia, expanded proximal fibula), orient the humerus to not be in posterior view, and get rid of the post-obturator notch in the ischium, which was later found to be erroneous.  But compare that to Peters' monstrosity.  The squamosal is just a triangular plate, lacking any posterior process so that the quadrate head is impossibly just floating freely posterior to it.  The quadrate has a narrow orbital/pterygoid process unlike anything but derived birds.  There are twenty-five presacral vertebrae unlike (?)all non-maniraptoran avepods, the caudals vary randomly in length by large degrees unlike any theropod, and the chevrons are all short as if it were a derived paravian.  In the pectoral girdle, there's some huge crescent where the sternum would be, but Sinosauropteryx lacks an ossified sternum.  The ilium is just wacky- tiny postacetabular process and concave dorsal margin.  The ischium is a thick blob with nary an obturator process.  Perhaps most sadly, Peters can't even draw the tibia as longer than the femur, which is (?)universal in small theropods, stated even in the crappy original Chinese description and is character 195 in Peters' analysis.

With his reconstructions being more or less fictional animals, it's no surprise the cladograms based on them will be equally unrealistic.  And lo, they are!  For fun, I give his cladogram from March 14th and applied phylogenetic nomenclature to see how it fares.  I listed the assumed position of taxa not yet included, as they are specifiers of various clades.  I also list where family-level taxa would have to be renamed due to ICZN rules.

Theropoda
|--Tawa
`--+--+--Staurikosaurus
   |  `--+--Segisaurus
   |     `--+--Guaibasaurus
   |        `--+--Marasuchus
   |           `--Procompsognathus
   `--Avepoda
      |--Coelophysoidea
      |  |--Dracoraptor
      |  `--+--Coelophysidae
      |     |  Coelophysis
      |     `--Dilophosauridae
      |        Dilophosaurus
      `--Neotheropoda (= Orionides, Avetheropoda, Neotetanurae)
         |--Ceratosauria (= Spinosauroidea, Carnosauria, Deinocheiridae)
         |  Megalosauria
         |  |--Spinosauridae (= Proceratosauridae)
         |  |  |--Proceratosaurus
         |  |  `--+--Deinocheirus
         |  |     `--+--Xiongguanlong
         |  |        `--+--Sinocalliopteryx
         |  |           `--+--Suchomimus
         |  |              `--Spinosaurus
         |  `--+--+--Dilong
         |     |  `--Guanlong
         |     `--Allosauroidea (= Neoceratosauria) (would need to be renamed Megalosauroidea)
         |        |--Abelisauroidea (= Metriacanthosauridae)
         |        |  |--Yutyrannus
         |        |  `--+--Sinraptor (assuming Metriacanthosaurus)
         |        |     `--Abelisauridae
         |        |        Majungasaurus (assuming Abelisaurus)
         |        `--Ceratosauridae (would need to be renamed Megalosauridae)
         |           Allosauria
         |           |--Allosauridae
         |           |  Allosaurus
         |           `--Megalosauridae (= Megalosauroidea, Carcharodontosauridae)
         |              |--+--Sinosaurus
         |              |  `--Monolophosaurus
         |              `--+--Acrocanthosaurus (assuming Carcharodontosaurus)
         |                 `--+--Eustreptospondylus (assuming Megalosaurus)
         |                    `--Ceratosaurus
         `--Coelurosauria (= Tetanurae)
            Maniraptoriformes (= Aviremigia, Tyrannoraptora, Bullatosauria)
            |--Compsognathidae (= Tyrannosauroidea, Arctometatarsalia, Microraptoria)
            |  |--Ornithomimosauria
            |  |  |--Compsognathus
            |  |  `--Struthiomimus (assuming Ornithomimus)
            |  `--+--+--Ornitholestes
            |     |  `--+--Microraptor
            |     |     `--Sinornithosaurus
            |     `--+--Fukuivenator
            |        `--+--Tianyuraptor
            |           `--+--Zhenyuanlong
            |              `--+--Alioramini
            |                 |  Alioramus
            |                 `--Tyrannosauridae
            |                    |--Gorgosaurus
            |                    `--Tyrannosauridae
            `--Maniraptora
               Pennaraptora (= Chuniaoae)
               |--Caenagnathiformes (= Oviraptoriformes)
               |  |--Therizinosauria
               |  |  |--Falcarius
               |  |  `--+--Jianchangosaurus (assuming Therizinosaurus)
               |  |     `--Rahonavis
               |  `--Oviraptorosauria
               |     |--+--Juravenator
               |     |  `--Sinosauropteryx
               |     `--+--Limusaurus
               |        `--Khaan (assuming Caenagnathus and Oviraptor)
               `--Paraves
                  |--Coeluridae
                  |  |--Tanycolagreus (assuming Coelurus)
                  |  `--Eotyrannus
                  `--Metornithes (= Eumaniraptora)
                     |--Dromaeosauridae (= Alvarezsauroidea, Alvarezsauria, Deinonychosauria)
                     |  |--+--Haplocheirus
                     |  |  `--Shuvuuia (assuming Alvarezsaurus)
                     |  `--+--Velociraptor (assuming Dromaeosaurus and Deinonychus)
                     |     `--Balaur
                     `--Avialae (= Averaptora)
                        |--Sinornithoides (assuming Troodon)
                        `--+--Jinfengopteryginae
                           |  Jinfengopteryx
                           `--+--Unenlagiinae
                              |  |--Anchiornis
                              |  `--+--Aurornis
                              |     `--Buitreraptor (assuming Unenlagia)
                              `--+--Sinovenator
                                 `--+--+--Eosinopteryx
                                    |  `--Xiaotingia
                                    `--+--Thermopolis "Archaeopteryx"
                                       `--Ornithothoraces (= Ornithes, Ornithopectae)
                                          |--Archaeopterygidae (= Saurornithes, Enantiornithes)
                                          |  |--Archaeornis
                                          |  `--+--+--Archaeopteryx
                                          |     |  `--Yixian embryo IVPP V14238
                                          |     `--+--Protopteryx
                                          |        `--+--Cathayornis (assuming Iberomesornis and Enantiornis)
                                          |           `--+--Pengornis
                                          |              `--Sulcavis
                                          `--Ornithurae
                                             |--Omnivoropterygiformes
                                             |  |--"Archaeopteryx" bavarica
                                             |  `--+--Mei
                                             |     `--Scansoriopterygidae
                                             |        |--Scansoriopteryx
                                             |        `--+--Yi
                                             |           `--+--Epidexipteryx
                                             |              `--Omnivoropteryx
                                             `--+--Jeholornis
                                                `--+--Jurapteryx
                                                   `--Pygostylia
                                                      |--Confuciusornithiformes
                                                      |  |--Wellnhoferia
                                                      |  `--Confuciusornis
                                                      `--Avebrevicauda
                                                         |--Chiappeavis
                                                         `--+--Sapeornis
                                                            `--+--Archaeornithura
                                                               `--+--Ichthyornis
                                                                  `--Aves
                                                                     |--Struthio
                                                                     `--Gallus (assuming Passer)


That's pretty funny.  Nomenclature fails in Allosauroidea and Compsognathidae, due to the weird topologies there where taxa not seen as closely related enough to need their neighbors as external specifiers suffer.  Ditto for dromaeosaurids vs. alvarezsaurs.  Also for archaeopterygids vs. enantiornithines, which no BAD analysis has ever recovered. 

As for the topology, there's that sister clade to avepods, whose oldest family name would be Procompsognathidae, though none of the members have phylogenetic definitions attached to them.  The division in neotheropods is sort of like Rauhut (2003), who had a huge Carnosauria.  The most basal megalosaurians are all actually basal tyrannosauroids.  Ornithomimids plus tyrannosaurids is a classic clade from Huene to Holtz, and Compsognathus being there reminds me of Olshevsky (1995) having it as a tyrannosaur.  Funny how Peters recovered Enigmosauria, though no doubt for completely different reasons than the dinosaur community, given the inclusion of compsognathids, Limusaurus and Rahonavis.  Troodontids and unenlagiines being avialans has been more popular recently (e.g. Agnolin and Novas, 2013), and the Archaeopteryx plus Enantiornithes pairing hearkens back to the BANDits' Sauriurae.  Similarly, Confuciusornis being closer to Aves than enants reminds me of Kurochkin's (2006) ideas.  And those are the only parallels I can make between Peters' non-consensus phylogeny and science.  My head's full of non-traditional phylogenetic proposals, but Deinocheirus as a spinosaur, or microraptorans as basal tyrannosaurs while Velociraptor is by alvarezsaurs?  That's just nuts.

So David, first I'd say you should start using the names Archaeornis, Jurapteryx and Wellnhoferia if you find these taxa away from the Archaeopteryx holotype.  But also, I think it would be amusing if you added the following- Megalosaurus, Hexing, Nothronychus/Erlikosaurus, Caudipteryx, Achillobator, Dromaeosaurus and Patagopteryx.

Peters' reconstruction of Archaeornithura meemannae from 3-26-16, preserved in case it's deleted like the first was (though Peters to his partial credit does say "Updated March 16, 2016 with new images. The beak, if present, is ephemeral, questionable. Only two scores changed.").

References- Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku Saizensen. 9, 92-119 (part 1); 10, 75-99 (part 2).
 
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.  

Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological Review. 86(suppl. 1), S45-S58.

Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer. 96 pp.

Fukuivenator thoughts

So we have another new theropod, Fukuivenator paradoxus (Azuma et al., 2016).  With Fukuiraptor, Fukuititan and Fukuisaurus also in existence, we just need a Fukuipelta, Fukuimimus and Fukuiornis to round out the Kitadani Formation.  This was first announced in an anonymous 2009 msn article (now defunct) and Shibata and Azuma's (2010) SVP abstract.  While those sources called it a dromaeosaurid, Azuma et al. interestingly say "it in fact represents a bizarre, basally branching maniraptoran theropod with a large number of autapomorphies."  Intriguing.  Let's check it out...

A partial skull is present (though the jugal, palatine, ?ectopterygoid and pterygoid are unillustrated) as well as a dentary fragment.  The weirdest thing is the gigantic premaxilla, which is so odd that I would question its identity and propose it's actually the right maxilla, except that it lacks the pneumatic fossa and foramina present on the left maxilla.  Contra the authors, the posterolateral process of the premaxilla is broken so (barring the presence of some really well preserved sutures on the maxilla) it cannot be determined if it extends posterior to the external naris like in dromaeosaurids.  Similarly, the supposedly large promaxillary fenestra (cited in the diagnosis as well) is actually small.  The text claims the lacrimal is T-shaped, but the figure shows the posterodorsal process is unpreserved, and the character is coded unknown in their matrix.  Although the authors claim the frontals have dromaeosaurid-like anterolateral notches and sigmoid supratemporal fossa edges, neither is visible in the figured left element.  Amusingly, Azuma et al. say one premaxillary tooth "exhibits a "ridge," on which denticles are absent."  Ah, you mean a carina? ;)  The other premaxillary tooth is indeed interesting in being D-shaped.  Surely the fenestra labeled "IX?" in figure 4  is the otic fenestra, and the basipterygoid processes are confusingly labeled as laterosphenoids twice in that figure.  The supposed medial eustachian foramina are the paired foramina of the basisphenoid recess.  Oddly, neither of the eustachian tube characters are coded, but the basisphenoid recess characters are coded correctly.  It's an interesting braincase, with no obvious dorsal or posterior tympanic recesses, a reduced basisphenoid recess and laterally diverging basipterygoid processes, most of which are troodontid-y.

Reconstructed skull of Fukuivenator paradoxus holotype, premaxilla and dentary flipped, and frontal shown in dorsal view (modified from Azuma et al., 2016).

An almost complete vertebral series is present.  The description states "ten cervical vertebrae are preserved, missing at least the atlas but the materials list only says "eight cervical vertebrae" are present.  Hmm.  The text states "the middle cervical vertebrae have a highly modified hyposphene-hypantrum with the hypantrum extending ventrally below the dorsal border of the neural canal (Fig. 6b)", but the figure shows nothing extending ventrally into the neural canal and I thought hyposphene-hypantrum articulations were absent from most theropod anterior dorsals, let alone cervicals.  "Dorsal centra are longer anteroposteriorly than tall dorsoventrally, unlike the dorsal centra of typical predatory theropods."  :|  Tell that to coelophysoids, compsognathid-grade taxa, troodontids, microraptorians...  "Likely pleurocoels are present in all dorsal vertebrae in the form of longitudinal fossae on the lateral surfaces of centra."  Haven't we progressed since the days of Osmolska et al. and Welles calling central fossae pleurocoels?  This is another character coded as dromaeosaurid-like, but the posterior dorsals (at least) actually lack pleurocoels as is standard for theropods.  Azuma et al. also state "the parapophyses of the dorsal vertebrae including the posterior ones are stalk-like as in derived alvarezsauroids and dromaeosaurids, though they are not as prominent as in the latter groups", but they're actually short.  My matrix uses a ratio of parapophyseal length (measured from the ventromedial corner to the apex) and centrum width, so that even Alxasaurus and some Sinraptor and Allosaurus vertebrae count as apomorphic (contra the unchanging TWG matrix), but Fukuivenator still falls short.  The authors state "the [sacral] zygapophyses are fused to each other to form a platform lateral to the neural spines, a feature also known in dromaeosaurids", but the figured sacrum lacks neural arches in the first four vertebrae.  Azuma et al. say the sacral ribs+diapophyses are "bifurcated distally to contact the ilium, a feature previously unreported in any other theropod", but this is true in e.g. the middle four sacrals of Gallimimus.  They also say "the most unusual feature is that the prezygapophyses of the middle caudal vertebrae are distally bifid (Fig. 6i), which has not been reported in any dinosaurs", but this is a standard dromaeosaurid character reported in e.g. Deinonychus (Ostrom, 1969) and Velociraptor (Norell and Makovicky, 1999).

Pes of Fukuiraptor paradoxus holotype, as shown in figure 7h on left, and with my reidentification of the phalanges on right (modified from Azuma et al., 2016).  Note the longer phalanges on digit II as in other theropods, and how digit II isn't particularly developed into a sickle-claw.

Contra the text and coding, the coracoid is proximodistally shallow, unlike pennaraptorans.  Surely, figure 7's caption is incorrect and the humeri shown are a right in anterior view and left in lateral view, not the left "in lateral (left) and posterior (right) view."  Also, the femur in figure 7f is in medial and posterior views, not lateral and posterior.  In figure 7h, the pedal phalanges are almost certainly placed incorrectly, with II-1 and II-2 switched with IV-1 and IV-2, which explains why digit II looks ridiculously short (see figure above).  "IV-1" has the standard shape of II-1 with the medial side highly concave (so it actually belongs on the other foot), plus I don't think any terrestrial Mesozoic theropod has IV-1 longer than II-1.  The measurement table is partly inconsistent as it has IV-2 subequal to IV-3 and IV-4 in length, unlike the figure.  Contra the text and coding, I don't think the second pedal digit looks particularly deinonychosaurian- Tanycolagreus has the same dorsally prominent distal articular surface on II-1, and the ungual in Ornitholestes is comparatively larger.  Alas, the text continues to use the II-III-IV manual digit identification, thanks to Xu, which at least Choiniere has disavowed (in his recent Ceratosaurus forelimb paper with Carrano).  It was an intriguing idea Xu, but it's long past time to let it die.

Azuma et al. add Fukuivenator to Turner et al.'s (2012) TWG analysis, which is unfortunately flawed by having numerous taxa left uncoded for entire sections of the matrix, and most of the codings retained from the original TWG matrix of Norell et al. (2001).  So don't put much credit into e.g. the joining of oviraptorosaurs and therizinosaurs, because the latter are only represented by the 2001 codings for Alxasaurus, Segnosaurus and Erlikosaurus.  No Beipiaosaurus, let alone Falcarius or Jianchangosaurus.  In any case, contra the text, they do not recover Fukuivenator as a maniraptoran.  In their trees, it's actually in a polytomy with compsognathids, Ornitholestes, ornithomimosaurs and maniraptorans.  They also didn't find "Anchiornis and Xiaotingia outside the Troodontidae", instead the base of Deinonychosauria is unresolved with those genera part of the polytomy.  Frustratingly, the authors never try excluding taxa a posteriori, so we don't know if e.g. the underlying structure of Turner's tree is still there and Fukuivenator can fall out in multiple places, or if it creates an actual polytomy between some taxa there.  Running the matrix myself, I find the latter is true- Fukuivenator's character combination creates a polytomy there even if it's pruned from the tree a posteriori.  Also, the Anchiornis+Xiaotingia situation exists because in a minority of trees these are sister to microraptorians+eudromaeosaurs instead of being basal troodontids.  Interestingly, the authors do check how parsimonious it is to place Fukuivenator in alternative positions and find it only takes three more steps to make it a paravian, deinonychosaur or dromaeosaurid. 

Adding Fukuivenator to the Lori matrix recovers it in a trichotomy with Ornithomimosauria and Maniraptora.  Makes sense considering its general basal coelurosaur grade anatomy.  There are a few dromaeosaurid-like characters- the fossa around the maxillary fenestra, the short ventral postorbital process, the squamosal shelf over its ventral process, the twisted paroccipital processes, the reduced crista prootica, the bifurcated caudal prezygapophyses.  But the vast majority are quite unlike paravians, and as seen above, many of the supposedly dromaeosaurid-like characters aren't present.  Andrea Cau recovered Fukuivenator as sister to Pennaraptora in his Megamatrix, which isn't far from my result as only alvarezsauroids and therizinosaurs are between the two Fukuivenator positions, basal members of which (e.g. Haplocheirus, Falcarius) are like Ornitholestes-grade coelurosaurs in much of their anatomy. 

References- Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp.

Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.

Anonymous, 2009. [3rd new species? Small-sized meat diet dinosaur to restoration Fukui] msn.com 3/18/2009 http://sankei.jp.msn.com/science/science/090318/scn0903182113002-n1.htm 

Shibata and Azuma, 2010. New dinosaurs from the Lower Cretaceous Kitadani Formation of the Tetori Group, Fukui, Central Japan. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 163A-164A.

Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.

Azuma, Xu, Shibata, Kawabe, Miyata and Imai, 2016. A bizarre theropod from the Early Cretaceous of Japan highlighting mosaic evolution among coelurosaurians. Scientific Reports. 6, 20478.

Sauropod Thoughts

So this month I've been concentrating on adding sauropods to the Database, going in reverse chronological order.  I find sauropods interesting because unlike theropods, their phylogenetics are almost completely unresolved.  I mean sure there's a basic outline and diplodocoids are pretty well researched, but there are huge swaths of cetiosaur-grade and titanosaur(iform/oid) grade taxa that can go almost anywhere between Vulcanodon and Neosauropoda on the one hand, and between Camarasaurus and Eutitanosauria on the other.  What follows are some thoughts on taxa or papers describing them...

Zby is based on mostly a forelimb that was originally assigned to Turiasaurus in a SVP abstract (Mateus, 2009), but later described as a new genus (Mateus et al., 2014).  But the seven characters listed by the latter authors as distinguishing the taxa are only determinable in their holotypes, not the referred specimens of Turiasaurus.  Given their stratigraphic and geographic proximity and the variation seen within theropod species, it seems plausible Mateus was right in 2009.

Holotype of Zby atlanticus (ML 368) (after Mateus et al., 2014).

I didn't realize how sad the descriptive situation is for Galeamopus. Sure the type skull was described almost a century ago, and the other skulls have featured in discussions of Diplodocus crania, but the postcrania of the type or the new Howe Quarry specimen (SMA 0011) have never been described.  We need not only a description of these, but also a paper describing all the supposed Diplodocus skulls, since according to Tschopp et al. (2015), no skulls can actually be referred to that genus.  Thus the Diplodocus OTUs in every other analysis are composites.

Despite having four generally stellar coauthors, the description of Abydosaurus (Chure et al., 2010) is... well... bad.  Even though it has seven supplementary documents, one of which is a spreadsheet of several hundred sauropodomorph tooth elongation indices, there's no measurement table for anything except two cervicals.  The most we get for the skull is "The four known skulls of Abydosaurus are nearly the same size, measuring approximately 0.5 m long and half as tall posteriorly."  You could have used less words and given us precise data for all four specimens.  And the table with the cervical measurements includes data for seven other sauropods, all but two of which are from previously published literature anyway.  Why spend space on data already out there instead of publishing new data?  The referred postcrania don't have repository numbers listed either.  Finally, the phylogenetic analysis goes out of its way to delete data from Wilson's original version, removing all but two non-neosauropods, combining genera to make four suprageneric OTUs, deleting characters that don't vary due to those removals, etc..  All this does is make it more tedious for future workers to add Abydosaurus to the base version of Wilson's matrix that tens of other papers use, and it's not like Wilson's 234 character, 27 taxon matrix is unwieldy for any 2010 computer to handle.  They just spent a lot of time and effort to make the analysis worse.

Proximal caudal vertebra of Normanniasaurus genceyi (MHNH-2013.2.1) (after Le Loeuff et al., 2013). Scale = 10 cm.

Normanniasaurus (Le Loeuff et al., 2013) is one of the most fragmentary taxa I've seen erected in the past few decades (the holotype consists of two presacral prezygapophyses, three fused sacral centra, an incomplete proximal caudal vertebra, an incomplete mid caudal vertebra, partial scapula, two ilial fragments, ischia [one partial, one fragmentary], a femoral fragment and a fibular fragment).  This is quite ironic considering Le Loeuff's famous 1993 paper declaring most European titanosaurs to be indeterminate (Macrurosaurus, Hypselosaurus, Aepisaurus, "Titanosaurus" lydekkeri).  Most of the supposedly diagnostic characters listed by the authors are either primitive for titanosauriforms (presacral vertebrae with hyposphene-hypantrum articulation; middle caudal vertebrae amphicoelous; ilium with anterolaterally expanded blade) or classic characters diagnosing larger clades within it (internal texture of presacral vertebrae camellate; proximal caudal vertebrae deeply procoelous; middle caudal vertebrae with anteriorly placed neural arch).  I'm very doubtful whether the remaining few caudal characters will prove diagnostic if anyone looks into them.  When added to Mannion et al.'s (2013) matrix, Normanniasaurus resolves as a eutitanosaur closer to Saltasaurus than opisthocoelicaudiines. 

Finally, Mannion and Otero (2012) did a superb job redescribing Argyrosaurus.  The taxon is only known from a forelimb, though apparently the entire skeleton was originally there but did not survive excavation.  This is my favorite kind of paper- redescribing a taxon known for decades in a modern context.  Mannion's one of my favorite sauropod authors, having redescribed Mongolosaurus (2010), "Brachiosaurus" atalaiensis (2013) and B? nougaredi (2013).  Mannion and Otero didn't include Argyrosaurus in a phylogenetic analysis, saying it has some plesiomorphic characters despite sharing many characters with derived titanosaurs.  I coded Argyrosaurus in Mannion et al.'s macronarian matrix, and I have to say the characters are well formed, so that coding is objective.  I'm not a sauropod expert, so I don't have the background comprehension of sauropod characters, but Mannion makes them easy to code.  Argyrosaurus ends up as a nemegtosaurid (sister to Mongolosaurus and Rapetosaurus), so despite Mannion et al.'s lack of many lithostrotians, the genus seems to be a member of the clade.  If it is a nemegtosaurid, Argyrosauridae Bonaparte 1987 has priority over Nemegtosauridae Upchurch 1995.  Interesting.  One critique of the paper is that it doesn't specify the horizon and locality of referred Argyrosaurus specimens (so I had to refer back to Powell 2003 and Bonaparte and Gasparini 1979), and proposes these are nomina dubia without detailed examination or comparison.

Will the next post get back to theropods? Depends on the papers published in the near future.  Come on, O'Connor, I dare ya. ;)

References- Mateus, 2009. The sauropod dinosaur Turiasaurus riodevensis in the Late Jurassic of Portugal. Journal of Vertebrate Paleontology. 29(3), 144A.

Chure, Britt, Whitlock and Wilson, 2010. First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition. Naturwissenschaften. 97(4), 379-391.

Mannion and Otero, 2012. A reappraisal of the Late Cretaceous Argentinean sauropod dinosaur Argyrosaurus superbus, with a description of a new titanosaur genus. Journal of Vertebrate Paleontology. 32(3), 614-638.

Le Loeuff, Suteethorn and Buffetaut, 2013. A new sauropod dinosaur from the Albian of Le Havre (Normandy, France). Oryctos. 10, 23-30.

Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society. 168(1), 98-206.

Mateus, Mannion and Upchurch, 2014. Zby atlanticus, a new turiasaurian sauropod (Dinosauria, Eusauropoda) from the Late Jurassic of Portugal. Journal of Vertebrate Paleontology. 34(3), 618-634.

Tschopp, Mateus and Benson, 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ. 3:e857.

Dracoraptor thoughts

So the Welsh coelophysoid has been published, with the dudebro name of Dracoraptor.  There seems to have been a lack of rigorous editing, as exemplified by "Podekosaurus", Eshanosaurus being a therizinosaurid (as opposed to -oid or -ian), BMNH and NHMUK being used in the same sentence for specimens, and that's just the first two pages.  I also think the kind of title this paper has is silly ("The oldest Jurassic dinosaur") since it just specifies position within a known range.  It's like having "The smallest dinosaur under two meters long" or "the westernmost dinosaur found in China." Pluses are it's in open access and the photos are color with great resolution.

As for the actual description, there's a discrepancy in the elements that are shown in the skeletal reconstruction (fig. 5), listed under the holotype, listed at the beginning of each anatomical area, and actually described.  For instance, figure 5 shows a squamosal, angular and articular as preserved (and not just as molds or tentatively identified), which are listed under holotype.  Yet only "more caudal elements of the lower jaws" are listed under the cranial elements heading, and none of these elements are described.  But a ?nasal is described and figured, but not in figure 5 or noted in the cranial intro.  In total it seems more written by committee than cohesive.

Color figures- holotype of Dracoraptor hanigani (NMW 2015.5G.3) with: A- supposed manual ungual III, B- supposed manual ungual II, C- supposed manual ungual I, D- supposed furcula (after Martill et al., 2016).  Black and white figures from left to right- pedal unguals I and II of Coelophysis bauri NMMNH P-42200 (after Rinehart et al., 2009); pedal ungual II in dorsal view of Liliensternus liliensterni syntype MB.R.2175 (after Huene, 1934); metacarpal II of Coelophysis? rhodesiensis holotype QG 1 (after Galton, 1971); furcula of kayentakatae holotype MNA V2623 in dorsal and anterior views (after Tykoski et al., 2002); ninth dorsal rib of Allosaurus fragilis USNM 8367 (after Gilmore, 1920). Red lines indicate element identifications I propose for Dracoraptor.

Note the maxillae are in medial view, as the figure caption states, but contra the text.  Contra the text and reconstruction, only the small anteriormost jugal groove probably articulated with the maxilla.  The longer ridge and groove on the main body is common in early dinosaurs (e.g. Herrerasaurus) and non-articulating.  The supraoccipital is in anterior (internal) view, not ventral view as stated by the text.  Note the large posttemporal fenestrae as in Silesaurus but unlike dinosaurs. 

The cervical is not opisthocoelous, contra the text, as the anterior central surface is slightly concave.  The supposed first caudal is near certainly a ?last sacral based on the broad transverse processes originating on the centrum (compare to e.g. Staurikosaurus).  The authors do call it "a partially sacralised element", but any ambiguity seems unnecessary.  The next element could easily be a sacral too, though its more fragmented condition makes this more uncertain.  I'm doubtful the supposed furcula is correctly identified.  One side is much narrower than the other, and each is curved in a different direction (thin side concave toward the outside of the angle).  Furcular arms are subequal in width, and those of coelophysoids (e.g. kayentakatae) are basically circular in section, so that twisting in Dracoraptor is not an excuse.  It's more probably a posterior dorsal rib, which are also similar in having a ridge along the outside corner. The tuberculum may be covered in matrix. 

Amusingly, Martill et al. state that on the humerus "a suture between the articulatory epiphyseal surfaces is well defined."  And for the ulna, "the proximal condyle surface is smooth and well defined, marked from the diaphysis by a prominent suture."  Dinosaurs aren't mammals, people.  They don't have separately ossifying epiphyses.  For the femur, "the femoral head (greater trochanter) is "hooked'."  Er, those are very different structures, not synonyms.  The authors say "A calcaneum is not present. Two distal tarsals (dt III & dt IV) and part of a putative third are present in a row."  I'm pretty sure no archosauriforms actually have three distal tarsals per pes (certainly no theropods do), so that 'putative third' is more likely the supposedly missing calcaneum, especially as it's placed right next to distal tarsal IV (confusingly labeled 'Ldtii'). 

The supposed "?Metaacarpal of digit I" [sic] is a metacarpal II, very similar to rhodesiensis, more elongate than metacarpal I and more robust than metacarpal III.  Even Peters got this right in his reinterpretation, which is especially sad for Martill et al..  While I haven't identified all of the phalanges in this block, it's clear Martill et al.'s statement "they are assumed to be from the left manus as they are associated with the left radius and ulna" is in error.  For instance, the phalanx underlying the proximal radius is too large to belong to any manual digit and is probably pedal phalanx III-1, while supposed manual unguals I and III lack flexor tubercles ("I" shows an obvious depression in that area) and at least "III" is virtually straight.  These unguals more nearly match pedal unguals of e.g. Coelophysis and Liliensternus, while supposed ungual II is manual due to its curvature and large flexor tubercle.  Among other phalanges, that at the distal end of metacarpal II matches a manual phalanx II-1, that on the proximal end of metacarpal II belongs to manual digit III, that between unguals "I" and "II" looks like its from pedal digit II, and the small one by the anterior end of the dorsal centrum would be manual IV-1.  The latter suggests a less reduced digit IV than coelophysids or Herrerasaurus.  Thus the manual reconstruction with its short penultimate phalanges and metacarpal ratios should not be trusted.

The phylogenetic analysis is  based on Nesbitt et al.'s Tawa matrix.  One good thing is that Martill et al. split Nesbitt's composite characters, which had the effect of Nesbitt assuming non-homology for e.g. subnarial foramina in different positions, jugal ridges of varying sharpness, etc..  A negative point is that Martill et al. do not order any characters "because it assumes a complete fossil record and a direction to evolution."  But ordering does not assume these, it merely observes that some states are more similar to others.  So e.g. if a taxon has 5 premaxillary teeth, it's more similar to a taxon with 6 teeth than to one with 3 teeth.  Ironically, some of Martill et al.'s states imply ordering, such as their "6+ premaxillary teeth" state, which only makes sense if taxa with 6 teeth are more similar to taxa with 7, 8, etc. teeth than to those with less than six teeth.  They recover Dracoraptor as the basalmost coelophysoid, sister to Coelophysidae. I like how they note "an unnamed clade containing Velociraptor, Allosaurus and Piatnitzkysaurus."  Ah, you mean that obscure group called Tetanurae. ;)

Luckily for me, I have a version of Nesbitt et al.'s matrix on hand with characters properly ordered, and basically every basal dinosauriform added (as used in my Chilesaurus post).  I've also made a ton of corrections, though have by no means gone through the whole thing yet.  I did go through Martill et al.'s Dracoraptor codings though, and 15% are miscoded.  These range from consequences of my reidentifications above, to general errors (e.g. while coded as lacking a promaxillary fenestra, you can't tell that as the maxillae are in medial view), to times where I don't think the authors understood the character (e.g. it's coded as having an anterior hollow on the astragalus, but as used by Nesbitt this is a feature that has been lost in all avemetatarsalians).  After correcting Dracoraptor's codings, it emerges as sister to Neotheropoda (Ceratosauria+Tetanurae; Averostra of some authors).  But Neotheropoda has an odd topology where Chilesaurus is sister to Velociraptor, then Ceratosaurus, Piatnitzkysaurus and Allosaurus form a clade.  Because this makes the basal condition of Neotheropoda very different than the consensus, I deleted the highly modified Velociraptor.  Recall from the Chilesaurus post that Velociraptor only clades with Piatnitzkysaurus and Allosaurus in Nesbitt's uncorrected trees because it's miscoded for 24% of the characters that make it seem more like a "normal" theropod.  After Velociraptor's deletion Neotheropoda has the standard topology, and Dracoraptor is sister to a Daemonosaurus+Chilesaurus clade, which is itself sister to Avepoda (Neotheropoda of some authors).  This is rather like Cau's result, where Dracoraptor grouped with Daemonosaurus and Tawa.  In my trees, Tawa is the next taxon stemward.  Because Chilesaurus' basal theropod status is my pet theory and the taxon is also highly modified, I tried deleting it as well.  The result then is that Dracoraptor forms a polytomy with Tawa, Daemonosaurus, Coelophysidae and more derived theropods.

Thus it seems Cau's and my matrices agree in placing Dracoraptor by Daemonosaurus.  Note both share three premaxillary teeth (also in Chilesaurus and Tawa), short snouts with few maxillary teeth (also in Chilesaurus, not in Tawa), a lacrimal flange that does not overlap the antorbital fenestra (not in Tawa; unknown in Chilesaurus), and elongate cervicals with single pleurocoels (also in Tawa; elongate centra but double pleurocoels in Chilesaurus). 

References- Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. Bulletin of the United States National Museum. 110, 1-154.

Huene, 1934. Ein neuer Coelurosaurier in der thüringischen Trias [A new coelurosaur in the Thuringian Trias]. Paläontologische Zeitschrift. 16(3/4), 145-170.

Galton, 1971. Manus movements of the coelurosaurian dinosaur Syntarsus and opposability of the theropod hallux. Arnoldia. 5(15), 1-8.

Tykoski, Forster, Rowe, Sampson and Munyikwa, 2002. A furcula in the coelophysid theropod Syntarsus. Journal of Vertebrate Paleontology. 22(3), 728-733.

Rinehart, Lucas, Heckert, Spielmann and Celeskey, 2009. The Paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History and Science Bulletin. 45, 260 pp.

Martill, Vidovic, Howells and Nudds, 2016. The oldest Jurassic dinosaur: A basal neotheropod from the Hettangian of Great Britain. PLoS ONE. 11(1), e0145713.

Nanotyrannus was a groovy baby

The title works best when read in Austin Powers' voice.

So a new paper's out, and let's just say it's not very good.  Schmerge and Rothschild's (2016) premise is that Nanotyrannus has a lateral dentary groove, so based on this and adding it to Brusatte et al.'s (2010) tyrannosauroid matrix, it's an albertosaurine instead of a tyrannosaurine, let alone a juvenile Tyrannosaurus.

The first problem is that the lateral dentary groove has never been a great character, since its depth varies so much between taxa that numerous genera have been coded both ways in different TWG analyses.  I would say for instance that the slight longitudinal depression in Dryptosaurus (Brusatte et al., 2011; fig. 3A) doesn't qualify, contra the authors.  Indeed, Carr (online) list additional tyrannosauroids he interprets as having the groove, which were not coded that way by Brusatte et al..  Of these, I'd agree with Raptorex, and the juvenile Bistahieversor and Tarbosaurus, but view e.g. the Bistahieversor type and Alioramus (altai's type) to be more like Dryptosaurus.  Thus I agree with Headden (DML, 2016) that the character shows ontogenetic variation.  But my main point is that it's so subjective that authors can't agree on which morphologies count, and that any attempt to quantify depth and continuity (many individuals have grooves that vary in depth along the dentary, most obviously visible in the CT scan of Tarbosaurus juvenile IGM 107/7) won't be easy.

Dentaries of tyrannosauroid taxa (some flipped) showing degree of lateral groove.  Left (top to bottom)- Bistahieversor juvenile paratype NMMNH P-25049 (after Carr and Williamson, 2010); Tarbosaurus juvenile IGM 107/7 (after Tsuihiji et al., 2011); Nanotyrannus BMRP 2002.4.1 'Jane' (after Schmerge and Rothschild, 2016); Raptorex juvenile holotype LH PV18 (after Sereno et al., 2009). Right (top to bottom)- Dryptosaurus holotype ANSP 9995 (after Brusatte et al., 2011); Bistahieversor holotype NMMNH P-27469 (after Carr and Williamson, 2010); Alioramus IGM 100/1844 (after Brusatte et al., 2012); Tyrannosaurus FMNH PR2081 (after Brochu, 2003).  I'd say those on the left have a strong groove while those on the right lack one, and the least Dryptosaurus has less of a groove than e.g. Bistahieversor or Tyrannosaurus, contra Schmerge and Rothschild's recoding of it.  Also note deeper grooves are common in juvenile tyrannosaurids.

The second problem is that even if Schmerge and Rothschild had an unassailable, quantifiable character difference between Nanotyrannus and Tyrannosaurus that was not known to be ontogenetically variable, it's STILL just one character.  They're being BANDit-esque in assuming one character can override the totality of character evidence.  Sure they go on to list five other characters supposedly shared between Nanotyrannus and albertosaurines, but not only are some obviously plesiomorphies  (maxillary fenestra placed posteriorly; rounder orbit; greater dentary tooth count), the authors actually got the maxillary fenestra character backwards (they claim "contact of the maxillary fenestra with the rostral margin of the antorbital fossa" unites Nanotyrannus and albertosaurines).  The juvenile Tarbosaurus and Maleevosaurus' type are sufficient to show it's expected in juvenile Tyrannosaurus anyway.  What about the cladistic analysis, you ask?  While I haven't seen the matrix yet*, I'm betting the authors coded Nanotyrannus as if it was an adult.  And we already know juvenile tyrannosaurines emerge more basally when coded as adults.  Carr (2005) found Nanotyrannus' holotype to be sister to Daspletosaurus+Tarbosaurus+Tyrannosaurus, and the Stygivenator holotype to be "in a polytomy consisting of Teratophoneus and all non-tyrannosaurine tyrannosauroids."  Tsuihiji et al. (2011) found their Tarbosaurus juvenile to emerge sister to albertosaurines+tyrannosaurines.  So this is an unsurprising result even assuming Schmerge and Rothschild's coding was competent.

Edit- the matrix is terribly undercoded.   Only one character of 27 coded for the nasal and lacrimal, the squamosal and quadratojugal are completely uncoded, the entire skull roof, palate and braincase uncoded, postdentary mandibular bones uncoded, only one postcranial character coded.  Did nobody check the matrix and notice the huge swath of question  marks?!  Seriously, Cretaceous Research, this is just sad.

Finally, the authors don't understand how to compute character changes, when they compare the steps necessary to move Nanotyrannus to Tyrannosaurus' sister OTU.  Note we're never presented with the actual number of steps such a move adds to the analysis, let alone told if that number is statistically significant.  Instead, Schmerge and Rothschild just show the distribution of their dentary groove character if the move is made.  Even accepting their coding is correct, the correct answer is that moving Nanotyrannus to Tyrannosaurus would add a single step, namely Nanotyrannus evolving that character in convergence with albertosaurines.  But the authors claim "this tree requires 5 more independent losses of the dentary groove than the tree proposed in this study."  Wha?!  How do they justify this?  Because for some never explained reason, the authors assume the groove is present along the entire 'backbone' of the cladogram, so that every tyrannosauroid that lacks the groove independently loses it.  How terribly unparsimonious!  This takes 11 steps in their tree 3B, whereas the most parsimonious phylogenetic reconstruction would take only four steps assuming the groove is present in the outgroup as the authors believe.  All that's necessary is the loss of the groove for taxa more derived than proceratosaurids, then independent development in Dryptosaurus, albertosaurines and Nanotyrannus.  I can only suppose the authors view the groove as being incapable of parallel development.  It's just formed by a branch of the (?)trigeminal nerve being more depressed into the dentary's lateral surface, hardly the "dramatic change (e.g., metamorphosis) [that] would need to be invoked to explain the loss of this feature through maturation" that Schmerge and Rothschild claim.

Cladogram of tyrannosauroids with deep lateral dentary grooves indicated with bold lines.  Middle- figure 3B from Schmerge and Rothschild (2016) showing their proposed evolution of dentary grooves.  Top- number of steps assumed by Schmerge and Rothschild, with steps indicated by circles.  Bottom- most parsimonious distribution of steps given an outgroup with dentary grooves.

In conclusion, I don't see how this paper made it to publication.  The anatomical structure in question has a controversial distribution, Tsuihiji et al. (2011) destroys the entire premise but is never addressed or referenced, the authors never even consider that coding Nanotyrannus as an adult is problematic (Carr 1999 showed that the holotype has immature bone grain, so even if it's an albertosaurine, it's a juvenile), and they don't understand how character state reconstruction works.  Using 'key characters', presumptions about reversability, an ignorance of the recent literature, misunderstanding cladistics... did Schmerge inherit BANDit-style biases working at the University of Kansas?

References- Carr, 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 19(3), 497-520.

Brochu, 2003. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memior. 7, 138 pp.

Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. PhD thesis. University of Toronto. 1170 pp.

Sereno, Tan, Brusatte, Kriegstein, Zhao and Cloward, 2009. Tyrannosaurid skeletal design first evolved at small body size. Science. 326(5951), 418-422.

Carr and Williamson, 2010. Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea. Journal of Vertebrate Paleontology. 30(1), 1-16. 

Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky and Xu, 2010. Tyrannosaur paleobiology: New research on ancient exemplar organisms. Science. 329, 1481-1485.

Brusatte, Benson and Norell, 2011. The anatomy of Dryptosaurus aquilunguis (Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American Museum Novitates. 3717, 53 pp.

Tsuihiji, Watabe, Tsogtbaatar, Tsubamoto, Barsbold, Suzuki, Lee, Ridgely, Kawahara and Witmer, 2011. Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. Journal of Vertebrate Paleontology. 31(3), 497-517.

Brusatte, Carr and Norell, 2012. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History. 366, 197 pp.

Carr, online 2016. http://tyrannosauroideacentral.blogspot.com/2016/01/by-way-that-groove-is-also-seen-in.html

Headden, DML 2016. http://dml.cmnh.org/2016Jan/msg00074.html

Schmerge and Rothschild, 2016. Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al., 1988. Cretaceous Research. 61, 26-33.

How not to redefine Ornithuromorpha

The answer is "any possible way."  As in, DON'T redefine Ornithuromorpha Chiappe et al. 1999, which has until now only been given two definitions, both node-based.  There's Chiappe's (2001) original "Patagopteryx + Vorona + Ornithurae" and his later (2002) more refined "Patagopteryx + Ornithurae", both based on the same topology, with a trichotomy between Patagopteryx, Vorona and Ornithurae.  Ornithurae in this case is Chiappe's version which was Hesperornis plus modern birds.

Yet in the description of their new bird Dingornis [edit] Dingavis longimaxilla (first mentioned in Wang et al., 2015), O'Connor et al. (2016) redefine Ornithuromorpha to be stem-based.  Their Ornithuromorpha is "The first ancestor of Neornithes that is not also an ancestor of the Enantiornithes, and all of its descendants."

Holotype of Dingavis longimaxilla IVPP V20284 (scale = 20 mm; after O'Connor et al., 2016).

First of all, will everyone PLEASE start paying attention to Phylocode Article 11.1- "All specifiers used in node-based and branch-based definitions of clade names, and one of the specifiers used in apomorphy-based definitions of clade names, are species or specimens."  Chiappe had the excuse that he was working over a decade ago.  But in 2016 that doesn't cut it.  And it's SO easy!  Just say "Passer domesticus < - Enantiornis leali".  Done!

Second, O'Connor et al.'s excuse is that "Although the proposed definition does not strictly equate with the published node-based definition, it does provide a formal definition for the current widespread usage of this term in most recent literature (Bell et al. 2010; O’Connor et al. 2010; S. Zhou et al. 2013a; M. Wang et al. 2015)."  Let's review those...
- Bell et al. (2010) in their description of Hollanda have Patagopteryx as the most basal taxon closer to modern birds than enantiornithines, so have no need for a different definition than Chiappe's and don't explicitly use one either.
- O'Connor et al. (2010) in their description of Longicrusavis have the same situation.
- Zhou et al. (2013) in their description of new Archaeorhynchus specimens don't even show a topology or mention Patagopteryx.  Notably, Zhou and Zhang (2006) in the original description of Archaeorhynchus just deleted Patagopteryx from the matrix they used without justification.  Why do you hate Patagopteryx so much, Zhou?
- Wang et al. (2015) in their description of new Gansus material do indeed place Ornithuromorpha in the wrong position, as they have Archaeorhynchus and Jianchangornis further from modern birds than Patagopteryx or Vorona, yet include them among Ornithuromorpha.  Note O'Connor was second author here.
So only one paper listed actually uses Ornithuromorpha as if it were a stem, and that shared a coauthor.  Hardly a convincing case that the usage is "widespread."  Sure there are other papers that use it this way, but almost all have O'Connor as a coauthor.  You don't get to use a name incorrectly tens of times and then say "looks like all the papers are using it this way, we better change it."

Third, by redefining names you're countering the entire point of phylogenetic nomenclature.  The beauty of the system is that regardless of your topology, you can apply clade names because their definitions stay constant.  Just find Passer and Ornithomimus in your tree and you'll always know where Maniraptora goes.  But if you pull a Sereno and declare Maniraptora to be "Oviraptor + Passer" instead, then your concept is no longer the same.  Ditto for Ornithuromorpha.

"Passer domesticus < - Enantiornis leali" is certainly a clade that deserves a name, but it's not the more exclusive Ornithuromorpha.  We didn't call the clade Ornithurae just because the BANDits got it wrong so often in the 90s, and we should aim to be better this time as well.



As an aside,  O'Connor et al. state "Although we do not consider [Gansus zheni] to be referable to Gansus, previously only known from the Xiagou Formation (You et al. 2006), we also recognize minor differences that suggest they are not referable to Iteravis huchzermeyeri."  After my in depth study, I'm skeptical.  Notably, O'Connor et al. did not add zheni to their matrix, although Iteravis emerged in a polytomy with Gansus. 
Is Dingavis from the same locality another synonym of Iteravis?  While it does seem to share the ventrally concave ischium with small mid dorsal process (also in Piscivoravis, Yanornis and Gansus), pedal digit IV is shorter (89% of III excluding unguals; compared to 99-110% in Iteravis), the ectethmoid (lacrimal of O'Connor et al.) is more acute, the carpometacarpus more elongated, and phalanx III-1 lacks the lateral process.  So offhand, I'd agree they're distinct.

References- Chiappe, 2001. Phylogenetic relationships among basal birds. In Gauthier and Gall (eds). New perspectives on the origin and early evolution of birds: Proceedings of the international symposium in honor of John H. Ostrom. Peabody Museum of Natural History. 125-139.

Chiappe, 2002. Basal bird phylogeny: Problems and solutions. In Chiappe and Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. University of California Press. 448-472.

Zhou and Zhang, 2006. A beaked basal ornithurine bird (Aves, Ornithurae) from the Lower Cretaceous of China. Zoologica Scripta. 35, 363-373.

Bell, Chiappe, Erickson, Suzuki, Watabe, Barsbold and Tsogtbaatar, 2010. Description and ecologic analysis of Hollanda luceria, a Late Cretaceous bird from the Gobi Desert (Mongolia). Cretaceous Research. 31(1), 16-26.

O’Connor, Gao and Chiappe, 2010. A new ornithuromorph (Aves: Ornithothoraces) bird from the Jehol Group indicative of higher-level diversity. Journal of Vertebrate Paleontology. 30(2), 311-321.

Zhou, Zhou and O'Connor, 2013. Anatomy of the basal ornithuromorph bird Archaeorhynchus spathula from the Early Cretaceous of Liaoning, China. Journal of Vertebrate Paleontology. 33(1), 141-152.

Wang, Clarke and Huang, 2015. Ornithurine bird from the Early Cretaceous of China provide new evidence for the timing and pattern of the evolution of avian skull. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 233.


Wang, O'Connor, Li and You, 2015. New information on postcranial skeleton of the Early Cretaceous Gansus yumenensis (Aves: Ornithuromorpha), Historical Biology. DOI: 10.1080/08912963.2015.1006217

O'Connor, Wang and Hu, 2016. A new ornithuromorph (Aves) with an elongate rostrum from the Jehol Biota, and the early evolution of rostralization in birds. Journal of Systematic Palaeontology. DOI: 10.1080/14772019.2015.1129518