Next up is Ornithomimosauria. The
Theropod Database has been updated with new ornithomimosaur info too. The published cladogram here is-
I included the Angeac taxon as an OTU, which had very few illustrated elements at the time of scoring- the cervical, tibia and distal metatarsus with a few other reported details like the toothless dentary. It emerged as an ornithomimosaur but since then good views of the known material have been made public and it's pretty obviously a ceratosaur similar to
Limusaurus or
Elaphrosaurus. This was always an easy possibility in the matrix, as it can move to Ceratosauria in the published matrix with only two steps. After rescoring it for the more complete remains, the Angeac taxon emerges in Ceratosauria sister to
Deltadromeus. Here's its new scorings
????????1? ?????????? ?????????? ?????????? ?????????? 0????????? ?????????? ???2??0??? ?????????- ????--???? ??0?0??10? ???{01}??0??0 000?1?1{01}0? ???????{01}01 ?01?02{01}0?{01} ?????????? ????????11 1000?0?110 1100??{12}?00 ?1?10?1000 1??0??001? ??0??000{01}0 ???001???? ???0?????? {01}?021??0?1 ?0?110???? ?000?0{012}0?? ??{01}0???1?? ?????0?002 ??1?0?0??0 ??100?0?0? ??????11{01}1 ?????????? 1??1??{23}??? ?????????? ????-22?00 1001??0??? ?????????? 0??0011??? ?2000????? ?0?????00? ?0?000??0? ????{12}???0? ????000??? ??0-?1???? 0?1?1{01}???? ??1-?????0 ?0?????0?? ??00010?0? ?00??????? 0???????00 ???011???? ??00?????? ??0???0??0 ??0??????? ?????????? ?????????? ????0??010 ?????????? ???000-?0? ?0?-01?00? ???1?0??{01}? ??0?1????? ?????????? ??000?-?1? 00???0?{01}?? ???000???- -??00-???? ??????0??0 0???????0?
When Angeac moves in the new matrix with added taxa, we get a little change where
Harpymimus is more basal and the polytomy is resolved-
Btw, I published a list of phylogenetic definitions in the supplementary material, including the first to define Ornithomimosauria on the type species- (
Ornithomimus velox < -
Tyrannosaurus rex, Shuvuuia
deserti, Therizinosaurus cheloniformis, Oviraptor philoceratops, Troodon formosus,
Passer domesticus).
Ours is one of the few published studies using the new
Deinocheirus skeletons, and unlike Lee et al. (2014) I recovered it basal to garudimimids and any other ornithomimosaurs except for
Hexing. It only takes 4 steps to move with other toothless ornithomimosaurs, so that's still quite possible. What seems implausible is it being a garudimimid, as that takes 14 steps. The supporting evidence for that in Lee et al.'s analysis was never great, and as shown in the
Sciurumimus section the scoring in Choiniere's analysis is pretty bad.
This is the first time
Hexing has been in an analysis besides the ornithomimosaur-only one in its original description, which did not include
Deinocheirus. It groups with
Deinocheirus, which is funny because of the size difference, but there would have to be something to fill up that ghost lineage if
Deinocheirus is so basal.
Shenzhousaurus and
Harpymimus are next.
Kinnareemimus takes three steps to move to Ornithomimosauria in the old matrix, but only one step in the new matrix. It emerges in a trichotomy with
Shenzhousaurus and
Harpymimus. An extra step moves it to Alvarezsauroidea, so that's still pretty uncertain.
 |
| Archaeornithomimus asiaticus paratype left femur in posterior view (AMNH 21800) (Courtesy AMNH). |
While I didn't recover
Deinocheirus by
Garudimimus, I did get
Beishanlong there as in Lee et al.. Also interesting is that
Archaeornithomimus groups there. That genus is known from a whole ton more specimens than suggested by Smith and Galton (1990), which I was able to examine at the AMNH. There are whole boxes of tibiae, and metatarsals, and phalanges, that could really use a new osteology. Where's
Archaeornithomimus bissektensis you ask? Its holotype femur went way too many places to include, lacking a unique combination of scores. Sues and Averianov (2015) did assign a lot of Bissekty material to the taxon, but I didn't include any of the Bissekty isolated composite taxa in case they're chimaerical. I still scored them though, so if anyone wants to experiment, the Bissekty Ornithomimosauria is-
???????0?? ?????????? ?????????? ?????????? ?????{01}0000 ?0???????? ?????????? ?????????? ?????????? ?????????? ??011010(01)0 ?011????0? 0?0??0???? ?????????? 10{01}0?2{01}00? ?01??0??{01}1 0??1?0???? ?0?0?0?00? ?????????? ?????01000 1100210000 1100000020 ?????????? ????0????? ???2?0?1?? 0???0????? ?00??0??0? ??00?????0 ???10??00? 1?1??????? ????0???0? ????????11 ?????????? ??010????? ??????0??? ?????2???? {01}???0?0?01 ??????10?? ???0?11??? ?210?????? 1??????00? 0????0?0?1 ????????0? ??1?000??? ?????0???? ??1??{01}001? ??1??00??0 10??????10 0(01)1000?0?0 ?????0???? 0???????0? ???1?????? ??0?0??0?? ??00?????? ?????????? ?????????? ?????????? ???00??010 ??0?-????? ??????-?00 ?00-010??1 -11100??00 ???????--? ???00?00?? 0?00?????? ?00?000100 0{01}0000???- -???{01}--??? ??-?0????? ?10?100??0
 |
| Archaeornithomimus? bissektensis holotype right femur (CCMGE 726/12457) (Courtesy of Averianov). |
The "Grusimimus" specimen GIN 960910KD ends up in Garudimimidae too, despite generally being seen as a potential juvenile
Harpymimus. Joining the two takes 4 steps, so is still quite possible. I wonder if this really is an endemic Asian clade. Interestingly,
Arkansaurus (which we were able to score based on the new description) is a basal garudimimid here, but it can move either a node outside the garudimimid plus ornithomimid clade, or to Ornithomimidae or to Tyrannosauroidea (where it ends up similar to where
Suskityrannus was placed by its authors) with one step. The Lori matrix isn't that great with pedal characters yet.
We also provide a definition for Ornithomimidae using
O. velox and not dependent on the controversial
Deinocheirus- (
Ornithomimus velox < -
Garudimimus brevipes). At the base of Ornithomimidae, we have
Nedcolbertia. While an ornithomimosaurian
Nedcolbertia's been recently proposed by Brownstein (2017) and Hunt and Quinn (2018), this is the first published analysis to find the result. That being said, only 3 steps move it outside Maniraptoriformes, where it falls out by
Zuolong and megaraptorans. Guess there was some real signal keeping it outside Tyrannoraptora in Dal Sasso and Maganuvo's (2011) TWiG analysis. Brusatte's redescription should shed light on
Nedcolbertia's anatomy, as should the description of BYU 19114 from the Cedar Mountain Formation, said to be similar by Hunt and Quinn.
Interestingly,
Ornithomimus velox does not group with
Dromiceiomimus (which includes
O. edmontonicus here), but is instead down by
Sinornithomimus. The only character really grouping traditional
Ornithomimus species together is metacarpal I being longer than metacarpal II, which was not a character in the Lori matrix but it still takes 3 steps to combine them. Note Claessens and Loewen (2015) in their excellent redescription just assume
Ornithomimus sensu lato is monophyletic based on the metacarpal length, and I don't think anyone's actually used
O. velox as its own OTU before this. It would be funny if
Sinornithomimus ended up actually being the Chinese sister taxon to
Ornithomimus.
Aepyornithomimus is sister to this pair in the published matrix, but in a trichotomy with
Ornithomimus,
Sinornithomimus and more derived taxa in the new version (not shown). It wasn't resolved in its original description's analysis using the Choiniere matrix, merely more derived than
Archaeornithomimus.
Next is a novel clade of
Tototlmimus and "
Gallimimus" "mongoliensis", the first time the latter has been included in an analysis. Only 3 steps are needed to move it sister to
Gallimimus bullatus, and
Tototlmimus follows.
Tototlmimus was poorly resolved in its initial analysis based on Kobayashi characters. Another new clade follows-
Rativates plus Kaiparowitz supposed
O. velox specimen MNA Pl.1762A plus
"Ornithomimus" sedens, the latter based only on the holotype. Again,
Rativates has only been analyzed once before in its initial description, using Choiniere's heavily misscored matrix, while
sedens and the Kaiparowitz specimen have never been analyzed before.
 |
| Ornithomimosaur coracoids in right lateral and proximal views. Top left- Beishanlong (after Makovicky et al., 2010). Bottom left- Anserimimus (after Barsbold, 1988). Top right- Sinornithomimus (after Kobayashi, 2004). Bottom right- Gallimimus (after Kobayashi, 2004). Peach dot indicates coracoid tubercle, green line indicates lateral edge of infraglenoid buttress. |
Here's probably a good place to say that ornithomimosaur phylogeny has suffered a similar fate to coelurosaur phylogeny lately because everyone reuses Kobayashi's characters and scores just like how everyone reuses TWiG characters and scores. So everyone gets an
Anserimimus plus
Gallimimus clade, then an American clade of
Struthiomimus and '
Ornithomimus'. But the former clade is only based on two coracoid characters, both of which are flawed. Above on the right we have Kobayashi's (2004) figure 88 from his thesis illustrating the characters. Top right is
Sinornithomimus and bottom right is
Gallimimus. The first character is "laterally offset infraglenoid buttress of the coracoid", represented by how much the green curve protrudes downward here. A bit more in
Gallimimus, but compare to the then unknown
Beishanlong coracoid in the upper left. It has a hugely protruding process but isn't a part of the
Anserimimus plus
Gallimimus clade. It's not a commonly shown perspective, but is also found in
Nqwebasaurus,
Allosaurus, etc.. The second character is "biceps tubercle positioned more anterior to base of posterior process", which is the peach dot in the figure. Here note that
Beishanlong also has this anteriorly positioned, but more problematically
Anserimimus in the lower left does not. Maybe the drawing's wrong, but the rest of that figure seems accurate (e.g. the manus) and detailed unlike some of the more schematic ones in Barsbold's works (e.g.
Adasaurus' pelvis). Note that the more recent Xu et al. (2011) ornithomimosaur analysis that finds the Kobayashi arrangment misscores
Beishanlong for both characters. Instead, the Lori analysis (and my previous unpublished TWiG analysis incorporating Kobayashi's and other ornithimomid-relevent characters) recovers a
Struthiomimus plus
Gallimimus clade and an
Anserimimus plus
Dromiceiomimus clade. Forcing my pairing of these four in Xu et al.'s matrix only adds 2 steps, but getting the standard Kobayashi arrangement of them in my matrix takes 8 steps, so here I think I really might be on to something.
 |
| Tyrannoraptoran femora in lateral and/or anterior views. Top left- Xiongguanlong (after Li et al., 2010). Top right- Alioramus (after Brusatte et al., 2012). Center- Timimus (after Benson et al., 2012). Bottom left- Archaeornithomimus (courtesy AMNH). Bottom right- Garudimimus in lateral view (after Kobayashi and Barsbold, 2005) and Gallimimus in anterior view (after Osmolska et al., 1972). Peach indicates accessory trochanter in anterior view, green outline indicates accessory trochanter in lateral view. |
Also in the
Struthiomimus plus
Gallimimus group, we get
Timimus as the sister taxon to the latter genus. But of course
Timimus was reassigned to Tyrannosauroidea by Benson et al. (2012). So what gives? Well, if you look at Benson et al.'s reasoning for rejecting an ornithomimosaurian identification, they say "The morphology of the accessory trochanter and the relatively anteroposteriorly narrow lesser trochanter of NMV P186303 are similar to those of derived tyrannosauroids such as
Xiongguanlong and tyrannosaurids. They are unlike the anteroposteriorly broad, ‘aliform’ lesser trochanter and prominent, triangular accessory trochanter of allosauroids, ornithomimosaurs ..." "NMV P186303 lacks several features present in all ornithomimosaurs, such as the ‘aliform’ lesser trochanter and prominent accessory trochanter. In contrast, the lesser trochanter of some tyrannosauroids is anteroposteriorly narrower, and the accessory trochanter forms a transversely thickened region, similar to the condition in
T. hermani (e.g.,
Tyrannosaurus)." "
T. hermani also possesses a proximomedially inclined (‘elevated’) femoral head, a synapomorphy of derived tyrannosauroids (e.g.,
Tyrannosaurus;
Xiongguanlong), that is absent in ornithomimosaurs."
First of all,
Gallimimus has an elevated femoral head too (Osmolska et al., 1972: Plate XLVII). Not as much as most tyrannosauroids', but neither does
Timimus. Of course tyrannosauroids have always been recognized as having aliform anterior trochanters as well, so this is a matter of degree. Their figure 19B does look anteroposteriorly narrower (~49% of total femoral width; ~49% in
Xiongguanlong, ~52% in
Alioramus), unlike
Archaeornithomimus (~69%). But
Garudimimus' ratio is ~50%. Also note figure 19C, also labeled as lateral view, looks broader (~64%) and no doubt had a slightly more anterior angle to the photo. Even ignoring
Garudimimus, something that depends so heavily on exact angle of perspective, especially considering taphonomy and how theropod femoral heads phylogenetically vary in their anterior angle compared to the distal end (basal forms are famously more anteromedially directed), is not great evidence in my opinion. What about that accessory trochanter? I agree
Timimus' is more tyrannosauroid in side view, but ironically because they're larger than at least
Archaeornithomimus and
Gallimimus (green highlight), contra Benson et al.'s statement. And again some ornithomimosaurs like
Garudimimus have large accoessory trochanters too. Regarding transverse width, I can't see a difference between e.g.
Alioramus and
Gallimimus above (peach highlight). I certainly wouldn't say
Alioramus' is thicker. So is
Timimus a tyrannosauroid or an ornithomimosaur? I don't think the evidence is great either way, and certainly no published analysis scores for these difficult to quantify degrees of trochanter size. Honestly, the biostratigraphy makes me think it will ultimately be some coelurosaur convergent with both. Maybe something like
Aniksosaurus, also Gondwanan Early Cretaceous with a tall and narrow anterior trochanter.
Finally,
Qiupalong joins the
Anserimimus plus
Dromiceiomimus clade. In its description, it grouped in the American clade, but that's the same Xu et al. (2011) analysis noted above that misscores
Beishanlong as lacking the supposed
Anserimimus plus
Gallimimus characters.
So that's the Ornithomimosauria. I think the Lori analysis does a good job here doing one of the things it's meant to- include a ton of taxa that have either never been analyzed or were only added singly and separately to existing analyses. Another point I like to emphasize is the hidden instability of our consensus. You might be thinking 'well your analysis seems very poorly supported if all of these tested changes only take 3 to 4 steps each'. Yet you can rearrange the entire tree of Xu et al.'s (2011) ornithomimosaur analysis to my topology and it just needs 5 more steps in total. And Brusatte et al.'s (2014) tree doesn't even find resolution between
Harpymimus,
Beishanlong,
Garudimimus,
Archaeornithomimus,
Sinornithomimus and the derived clade. Overall I'd say this is the best ornithomimosaur analysis published, in taxon number, character number and robusticity of results.
Next, alvarezsauroids...
References- Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur,
Gallimimus
bullatus n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia.
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Barsbold, 1988. A new Late Cretaceous ornithomimid from the
Mongolia People’s Republic. Journal of Paleontology. 1988(1), 122-125.
Smith and Galton, 1990. Osteology of
Archaeornithomimus asiaticus (Upper
Cretaceous, Iren Dabasu Formation, People's Republic of China). Journal of Vertebrate
Paleontology. 10(2), 255-265.
Kobayashi, 2004. Asian ornithomimosaurs. PhD thesis. Southern Methodist University.
340 pp.
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Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521.
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