tag:blogger.com,1999:blog-32484128038147302502024-03-17T01:49:02.127-07:00The Theropod Database BlogHere's a place where I can post my thoughts on new papers, provide updates on my projects, and post info that will eventually be on my website The Theropod Database - <a href="http://theropoddatabase.com/">http://theropoddatabase.com/</a> . It will center on theropods, but may delve into other topics as well such as phylogenetics.Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.comBlogger264125tag:blogger.com,1999:blog-3248412803814730250.post-79800347046076952602023-12-31T11:34:00.000-08:002023-12-31T11:34:42.849-08:00The Avian Acetabulum: Feduccia grasping at straws<p>It's not every day that we get an article arguing against the dinosaurian origin of birds any more, with that consensus so strongly supported and many of its detractors being deceased. But last week Feduccia published "The Avian Acetabulum: Small Structure, but Rich with Illumination and Questions", so in the spirit of the Dinosaur Mailing List of the early 2000s, let's have a fun critique of it.</p><p>Feduccia's attempted main point is an exercise in what Makovicky and Dyke (2001) described as naive falsification over twenty years ago - taking some feature shared by birds and dinosaurs and claiming it's not homologous in those groups, and via a mysterious step 2 concluding phylogeny is in shambles and birds must be non-dinosaurian. Feduccia's (2023) character de jure is the open acetabulum, classic dinosaurian character still seen in birds, and we can use <i>Silesaurus </i>to demonstrate Makovicky and Dyke's entire point that makes Feduccia's argument invalid and useless even if he was correct about the anatomy.</p><p>Traditionally an open acetabulum supports dinosaurian monophyly, but I'm a fan of the recent idea that silesaurs are a basal grade of ornithischian and it turns out <b>dun dun DUN</b> silesaurs have closed acetabula. Thus genasaurs and saurischians most parsimoniously evolved open acetabula convergently. According to Feduccia's logic, there goes the idea that ornithischians are closely related to saurischians and archosaurian phylogeny must be reevaulated. But actually no, the vast majority of evidence still strongly supports ornithischians being closer to saurischians than pterosaurs, aphanosaurs, suchians, phytosaurs, etc. because it's not like this example of convergence magically creates evidence ornithischians are related to something else. That would be naive falsification.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi7rdgJ9NzFIL1RrmWtlcVqM1k4p7iUyr9wTOx-UNOz09cCVcetRR_L6gCaRbgYT5Z6mfHkUKV26X2SPOhIatdEmcgxGhjNy08RPKbDBPNPlg91XVSqwb0doC7mwaA_7QhSU3E1EFntnAd6f3N-nmZXDWDvNyAyGylHH_6W2oSkG1oHL-FmVVDFWrWFMgE/s965/PostosuchusFeduccia.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="648" data-original-width="965" height="215" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi7rdgJ9NzFIL1RrmWtlcVqM1k4p7iUyr9wTOx-UNOz09cCVcetRR_L6gCaRbgYT5Z6mfHkUKV26X2SPOhIatdEmcgxGhjNy08RPKbDBPNPlg91XVSqwb0doC7mwaA_7QhSU3E1EFntnAd6f3N-nmZXDWDvNyAyGylHH_6W2oSkG1oHL-FmVVDFWrWFMgE/s320/PostosuchusFeduccia.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Top- Figure 3a of Feduccia (2023) illustrating (left to right) <i>Stegosaurus</i>, <i>Allosaurus </i>and '<i>Postosuchus</i>'. Bottom- actual <i>Postosuchus </i>pelvis after Weinbaum (2002).<br /></td></tr></tbody></table><br /></p><p>So anyway, did you know "Late Triassic “rauisuchians” like <i>Postosuchus </i>exhibit an entirely theropodan pelvic anatomy, except for the acetabulum, which is largely closed."? Because I didn't. The first thing that's funny is that Feduccia's figure of <i>Postosuchus</i>' pelvis (Fig. 3a) is copied from Figure 15 of Chatterjee's 1985 original description of the genus, and we've known at least since Long and Murry 1995 (TWENTY-EIGHT YEARS AGO) that it's a chimaera using the ilium of <i>Lythrosuchus </i>and the pubis of <i>Shuvosaurus</i>. It's details like this that demonstrate how Feduccia et al. are stuck in the past. In any case, <i>Postosuchus </i>has some very non-theropodish pelvic features like the low ilium, huge supracetabular buttress, only two sacral attachments, non-contacting pubis and ischium, absent obturator flange and the characteristic 'rauisuchian' medioventral tilt, but Feduccia doesn't know this because he's stuck in Chatterjee's 1980s world where <i>Postosuchus </i>might as well be a tyrannosaur ancestor. <br /></p><p>Right away we get "An open acetabulum is considered one of the most unchallengeable synapomorphies of dinosaurs and birds as opposed to stem dinosauromorphs and dinosauriforms that had not yet achieved fully upright posture and still exhibit a closed or partially closed acetabulum", but as shown by potentially ornithischian silesaurs, nope. You can go all the way back to Ferigolo and Langer (2007) for the concept, recovered in phylogenetic analyses at least by 2016 by Cabreira et al. and more recently and explicitly argued by Müller and Garcia (2020). The latter don't even mention the word "acetabulum" because modern phylogenetics has long since moved past "unchallengeable" key characteristics.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4gHeklDbZO6PNGL_gGouk8C5Yxqf7MttfKtMYNxXZs29Rss9Hpb9RwPce2ZQ-urfEcEvgVKqbVVTgwo5iSUNx7x2jpMGtCH_UdkLfUS-5WsXeB4oov6XG9WSHZJ0pxoztGh5iYuq34WSjrDjN3CFYSNRYmr1X2C4bu6e33vjcXXW5Qe4-He5spjYYIAc/s797/NovasPelvisFig.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="325" data-original-width="797" height="130" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4gHeklDbZO6PNGL_gGouk8C5Yxqf7MttfKtMYNxXZs29Rss9Hpb9RwPce2ZQ-urfEcEvgVKqbVVTgwo5iSUNx7x2jpMGtCH_UdkLfUS-5WsXeB4oov6XG9WSHZJ0pxoztGh5iYuq34WSjrDjN3CFYSNRYmr1X2C4bu6e33vjcXXW5Qe4-He5spjYYIAc/s320/NovasPelvisFig.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Figure 2 of Novas (1996).<br /></td></tr></tbody></table> </p><p>Most of Feduccia's paper seems to be written in response to a straw man BAD (Birds Are Dinosaurs) scientist imagining "fully open acetabulum" is to be taken literally, having no medial enclosure at all, and don't you know it, that's often untrue in birds and other pennaraptorans. In reality, an "open acetabulum" in Dinosauria is contrasted with having only a narrow slit as in <i>Lagosuchus</i>, <i>Ornithosuchus </i>or indeed as illustrated in Chatterjee's '<i>Postosuchus</i>' chimaera. Just look at Figure 2 of Novas' classic 1996 work on dinosaur monophyly which shows the dinosaurian perforate acetabulum represented by <i>Herrerasaurus </i>with medial enclosure around the edges just as Feduccia illustrates in so many birds. Even Feduccia's own figure of <i>Stegosaurus</i>' pelvis (which like his <i>Allosaurus </i>pelvis is ultimately copied from Marsh's papers from the 1800s- such a contemporary reference!) meant to show the "open acetabulum in dinosaurs" has a significant medial rim. </p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhMLCWPM7514g8kdgKcaTcJzrtsAjwSv6iOHMYQOXnYYTt6QcZh8eKf1TKScKjXVAUnlnyNDR5OFXExFHzt9iChsLG5xXFx9exkPiJEACil5_dWL_PvCpdPepndR-qc_C0LYAd-BEh5J2QulT8NTt4KNWz_c7nCt9HUrahH5Ut7gxA5a4Xvj09IaFlLj5M/s1440/MeleagrisPelvis.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1440" data-original-width="891" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhMLCWPM7514g8kdgKcaTcJzrtsAjwSv6iOHMYQOXnYYTt6QcZh8eKf1TKScKjXVAUnlnyNDR5OFXExFHzt9iChsLG5xXFx9exkPiJEACil5_dWL_PvCpdPepndR-qc_C0LYAd-BEh5J2QulT8NTt4KNWz_c7nCt9HUrahH5Ut7gxA5a4Xvj09IaFlLj5M/s320/MeleagrisPelvis.png" width="198" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Meleagris </i>pelvis (top) as redrawn in Baumel and Witmer (1993) from (middle) Harvey et al. (1968), and (bottom) different specimen from Butendiek (1980).<br /></td></tr></tbody></table><br /><p>Amusingly, Feduccia says "The Nomina Anatomica Avium, like most modern avian anatomical references, portrays <i>Gallus </i>(domestic chicken) and <i>Anas </i>(domestic duck) with fully open, dinosaurian acetabula, unlike older images (Figure 1), which show substantial medial acetabular walls." His Figure 1 has drawings of "<i>Gallus</i>, Encyclopaedia Brittanica, 1911; <i>Anas</i>, The Vertebrate Skeleton, Cambridge Press, 1897", which in addition to being downright archaic, are a pop encyclopedia and general vertebrate anatomy book, neither of which seem like somewhere to trust little anatomical details from. Also, the Nomina Anatomica Avium doesn't even figure the pelvis of <i>Gallus </i>or <i>Anas</i>, it has <i>Meleagris</i> (Figure 4.15). And that is credited as being redrawn from Harvey et al.'s (1968) <i>Meleagris </i>osteology, which indeed shows a larger acetabular foramen than Feduccia's 1911 <i>Gallus </i>or 1897 <i>Anas</i>. But maybe it's a fluke. Let's check Butendieck's 1980 <i>Meleagris </i>osteology and nope, the foramen is even larger there. Guess it's not a modern cladist conspiracy.</p><p>Feduccia insists on staying behind the times when he writes "The continued illogical application of “phylogenetic nomenclature” [70], “phylonyms” [9], and the arbitrary redefinition of established taxon names necessitates the following nomenclatural clarifications. ... “Archosauria” is also used <i>sensu traditum</i> [71,72] and is therefore equivalent to the Archosauriformes of Nesbitt [73], Ezcurra [74], and de Queiroz et al. [9]" and every other work dealing with archosaur phylogeny and nomenclature since the 90s! Oh, except Benton's (1999) <i>Scleromochlus </i>redescription he cites as reference 72, which proposed the name Avesuchia for the crown group which nobody ever used again. Why Feduccia insists on using a concept outdated by thirty-some odd years and expects to be taken seriously is a mystery. Oh, and in his Table 1 Feduccia defines Ornithurae as "All those birds are more closely related to extant birds (Neornithes) than they are to Enantiornithes", so look who's applying phylogenetic nomenclature now. Guess it <b>can </b>be logical as a concept, who would have guessed?</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjRi_r9KWSadE517hrDz6d-Dapvhh2rm0IAqjuQHn2XrkT6Nv3o1ddLiEQInMQr6ytLkH_MRARJqUeb4n3sj1XiNJf1KQwJ17MCctf0QhLEoEYgmwt2rWdEN4v-HTqlBshtirLk6CI7ZAVzNuo-k_o0HVLlOAJs-yQc7LiAS__9EMXcwNPJCsOeD1mT3R8/s890/FeducciaAntitrochanter.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="754" data-original-width="890" height="271" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjRi_r9KWSadE517hrDz6d-Dapvhh2rm0IAqjuQHn2XrkT6Nv3o1ddLiEQInMQr6ytLkH_MRARJqUeb4n3sj1XiNJf1KQwJ17MCctf0QhLEoEYgmwt2rWdEN4v-HTqlBshtirLk6CI7ZAVzNuo-k_o0HVLlOAJs-yQc7LiAS__9EMXcwNPJCsOeD1mT3R8/s320/FeducciaAntitrochanter.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Top left- unfused <i>Meleagris </i>pelvis showing iliac contribution to the acetabulum (25) is larger than ischial contribution (64) (after Butendieck, 1980). Top right- <i>Sinornis </i>pelvis showing antritrochanter (vant) almost entirely on the ilium (after Sereno et al., 2002). Bottom- <i>Buitreraptor </i>(left) and Berlin <i>Archaeopteryx </i>(right) showing antitrochanters (at) (after Agnolin and Novas, 2011).<br /></td></tr></tbody></table><br /></p><p>It's not just acetabular closure that Feduccia is examining, he also comments on the distribution of supracetabular crests and antitrochanters. For the latter, he notes "A structure homologized with the avian antitrochanter has been reported in ... a miscellany of coelurosaurs [16,102,103]. There is no osteological evidence that these structures are homologous: the avian antitrochanter forms primarily<br />[69] or almost exclusively [90] from the ischium, whereas the so-called “antitrochanters” of these nonavian taxa are almost exclusively iliac in composition ... Moreover, while continuity of function is not a requirement of homology, it should be noted that putative “antitrochanters” of so great a range of taxa cannot possibly have performed the specialized mechanical role that the true antitrochanter does in extant birds as part of their peculiar hindlimb locomotory system [90,104]." So the antitrochanters of coelurosaurs aren't actually antitrochanters because they're on the ilium! But wait, let's pull out Butendieck's turkey osteology again, and oh no! <i>Meleagris </i>has an antitrochanter mostly formed by the ilium! And while Feduccia says "In both enantiornithines and ornithurines, the maintenance of balance in obligately bipedal locomotion is linked to the presence of an antitrochanter", Sereno et al. (2002; in a figure Feduccia specifically cites) shows that in the enantiornithine <i>Sinornis </i>the antitrochanter is almost entirely on the ilium despite supposedly having the same function as in living birds! That's also true in the Lecho Formation enantiornithine pelvis PVL 4042 by the way (Walker and Dyke, 2010: Fig. 14). Given even those three examples, when Feduccia says "The putative “antitrochanters” of alvarezsaurs and therizinosauroids are iliac in composition", or that in <i>Rahonavis </i>"the purported antitrochanter, however, is iliac in origin", it lacks any force of argument whatsoever. Since it CAN be that way in birds and even supposedly have the same function, it can also be that way in bird ancestors and relatives.</p><p>Similarly, for <i>Archaeopteryx </i>Feduccia claims "Agnolin and Novas [115] (Figure 3B) interpreted a slight swelling on the posteroventral rim of the acetabulum in the Berlin specimen as an antitrochanter, but this interpretation seems incorrect because (1) it is positioned on the posteroventral rim of the acetabulum, whereas the avian antitrochanter is posterodorsal to the acetabulum [69,90,104]; it is composed solely of the ilium, whereas the avian antitrochanter is composed exclusively or principally from the ischium." But as the ischial peduncle doesn't wrap beneath the acetabulum, the structure is clearly on the posterodorsal rim, and is furthermore basically identical in structure and position to that of <i>Sinornis </i>(and <i>Buitreraptor </i>shown next to it, contra Feduccia's claim unenlagiids lack antitrochanters) except for being a bit more slender than in <i>Sinornis</i>. How Feduccia can have looked at these figures and think <i>Sinornis</i> has an antitrochanter but <i>Archaeopteryx </i>does not is beyond me. Another example of Feduccia's confusion is when he states "<i>Avimimus portensosus</i> [sic], however, possesses an ischial antitrochanter that, therefore, appears homologous with that in birds" but then notes <i>A. nemegtensis</i> was diagnosed in part by Funston et al. (2018) as lacking an antitrochanter. Yet Funston et al.'s same figure cited by Feduccia shows the antitrochanter in <i>A. portentosus</i> is largely on the ilium (as Figure 4G doesn't even show the ischium) and indeed, <i>A. nemegtensis</i> doesn't preserve a proximal ischium at all so if the antitrochanter were ischial neither Funston et al. nor Feduccia would even know it lacked one! The basic conclusion is Feduccia doesn't know what he's looking at, making his attempted review of antitrochanter distribution useless.<br /></p><p>Feduccia is no better when discussing supracetabular crests, as evidenced when he says "Although Paul (2002) claims that a supracetabular crest is present in archaeopterygids, it is absent in the London, Berlin, Eichstatt and Munich specimens (Figure 8d)." But Paul's Plate 24Ab clearly shows a convex outer rim in ventral view that defines a supracetabular crest in a photo of a cast of the London specimen, whereas Feduccia's Figure 8d is a horizontally oblique drawing of a reconstructed generic <i>Archaeopteryx </i>pelvis that couldn't show lateral convexity in dorsal/ventral view even if it was there. So far from countering Paul, the medium is inferior, the subject is inferior, and the perspective would make evaluation impossible in any case. Indeed, Paul's entire Plate 24 is an effective argument against everything Feduccia claims about antitrochanters and supracetabular crests, showing for example that tyrannosaurids lack a supracetabular crest, so apparently it wasn't integral to a functioning theropod gait, and that young <i>Meleagris </i>lacks an ossified antitrochanter, so maybe at least some of these theropods with seemingly small or absent antitrochanters had larger cartilaginous structures that enabled them to be functionally similar to living birds and/or <i>Avimimus portentosus</i>.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiW6l73W6nXglbvwRhpf4xPHNF_LVJ71TBBLRsOD24XHvJ0l34VzB5s5d3jfUIh_9orIjdyHbVMyQJi9jDk7nScQUAVXbPG6va_h3_lby5spBCWb0oBmGdtRXrE8JU4jfE77ddGzm1FYOJD1fLjhzzv9v6ERxB8HVqzwoATSW_Rn9adfsQpZTkQkFHCbss/s1151/FeducciaCoelophysis.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1095" data-original-width="1151" height="304" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiW6l73W6nXglbvwRhpf4xPHNF_LVJ71TBBLRsOD24XHvJ0l34VzB5s5d3jfUIh_9orIjdyHbVMyQJi9jDk7nScQUAVXbPG6va_h3_lby5spBCWb0oBmGdtRXrE8JU4jfE77ddGzm1FYOJD1fLjhzzv9v6ERxB8HVqzwoATSW_Rn9adfsQpZTkQkFHCbss/s320/FeducciaCoelophysis.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Right- Feduccia's (2023) Figure 5 (caption below). Top left- <i>Megapnosaurus rhodesiensis</i> pelvis (after Raath, 1977- see, I can cite old papers too!). Bottom left- <i>Oksoko </i>pelvis (flipped horizontally, after Funston et al., 2020).<br /></td></tr></tbody></table><br /><p>My favorite part of this paper though is Figure 5, which shows drawings of the "Pelvis of the basal theropod <i>Coelophysis </i>(a) compared to an array of primarily ground-dwelling birds: (b) Solitary Tinamou (<i>Tinamus solitarius</i>), (c) Wild Turkey (<i>Meleagris gallopavo</i>), (d) Japanese Quail (<i>Coturnix japonica</i>), (e) Greater Roadrunner (<i>Geococcyx californianus</i>), (f) Emu (<i>Dromaius novaehollandiae</i>), (g) Bush Moa (<i>Anomalopteryx didiformis</i>), and (h) Elephant Bird (<i>Aepyornis hildebrandti</i>)." The first thing of note is that the <i>Coelophysis </i>drawing is highly inaccurate, being a copy of Colbert's (1989) illustrations by Lois Darling, which "do not depict the anatomy of <i>Coelophysis </i>accurately" (Downs, 2000). There are no pubic or obturator foramina, no obturator plate on the ischium, the supracetabular crest is not confluent with the ventral postacetabular edge, and rather importantly for this paper's subject the ilium is lacking the medial wall on the dorsal part of the acetabulam (that is partially ovelapped laterally by the supracetabular crest) and the large antitrochanteric surface formed by the ischial peduncle and ischium. While I agree with Feduccia the latter structure is not homologous to the maniraptoran antitrochanter (which is on the outside rim of the acetabulum), it does combine with the dorsal section to make a significant surface medial to the acetabular rim. Again, Feduccia is using a 30+ year old source that is well known to be wrong.</p><p>And what's even the point of the figure? "Note the contrast between the condition in these cursorial birds, which presumably should most closely approach the putatively ancestral theropod condition with respect to their acetabular morphology, and the condition in <i>Coelophysis</i>." First of all, why should an avian/neornithine anatomy that evolved in the Late Cretaceous converge with or regress to a Triassic group's morphology? Second, modern birds have a functionally different gait from non-maniraptoriform theropods like coelophysoids at least, where the femur is more horizontal and less mobile, so even in terrestrial birds I don't see why any special acetabular anatomy would be similar. Third, Feduccia thinks oviraptorosaurs are birds (e.g. Feduccia and Czerkas' 2015 "Testing the neoflightless hypothesis: propatagium reveals flying ancestry of oviraptorosaurs") and oviraptorids like <i>Oksoko </i>happened to have an even more open acetabulum than Darling's fake <i>Coelophysis </i>and a pelvis much more superficially similar to coelophysoids than to modern birds. So according to Feduccia, some cursorial birds DID approach what he imagined the ancestral theropod condition to be like. He just chose to only figure cursorial <b>crown </b>birds, which aren't very similar to coelophysoids in pelvic structure.<br /></p><p>Looks like my work in the Lori description (Feduccia's reference 52) got cited! "Reliance on simple tabulations of step count difference between constrained and unconstrained trees [52] to determine whether the most basal members of pennaraptoran clades have been correctly resolved by parsimony analysis of current data matrices is naive and takes no account of the statistical significance of differences in tree populations [50]." Oh noes! I was naive! And what's this reference 50 I should have taken into account? <a href="http://theropoddatabase.blogspot.com/2015/01/bandit-cladogram-evaluated-james-and.html">James and Pourtless 2009</a>! The study that actually found birds as maniraptorans, coelurosaurs, theropods and dinosaurs, but which hid it by only figuring majority rule bootstrap trees and always pruned out <i>Effigia </i>because it falling out as an ornithomimosaur was too embarrassing of a result even for them! And that was even after scoring all dinosaur manual characters as unknown, even in the pentadactyl <i>Eoraptor </i>and <i>Herrerasaurus </i>with zero controversy of digital homology! Bwa ha ha! Yeah, those are some methodologies I really should have taken to heart. Hey, at least Feduccia <a href="http://theropoddatabase.blogspot.com/2012/01/almost-famous-im-misquoted-in-feduccias.html">cited me as the correct author this time</a>!</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhzFVptqPtpGcfc0KQA4MtNqA2CiHJPLpHWMPANPZMBz3zkWe8SG0SXNIwZgeaNN0DP9wTp3u8VYbsc61yMS_Womw5guiPFTx7zPxSqCPHU0jnJkgHs5yXVBl62LlE3ClHvh6M2VLOg42Ocm5tzBorzbRT93cgedFaKVNdA9C934XigzlNefUeQuEcdizw/s1012/FeducciaIlium.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1012" data-original-width="569" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhzFVptqPtpGcfc0KQA4MtNqA2CiHJPLpHWMPANPZMBz3zkWe8SG0SXNIwZgeaNN0DP9wTp3u8VYbsc61yMS_Womw5guiPFTx7zPxSqCPHU0jnJkgHs5yXVBl62LlE3ClHvh6M2VLOg42Ocm5tzBorzbRT93cgedFaKVNdA9C934XigzlNefUeQuEcdizw/s320/FeducciaIlium.png" width="180" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Top- <i>Ornitholestes </i>holotype ilium and proximal pubis (after <a href="http://research.amnh.org/paleontology/search.php?action=image_detail&specimen_id=147302#">AMNH's 2007 digital collections</a>). Middle- <i>Allosaurus </i>USNM 4734 ilium (after Gilmore, 1920). Bottom- <i>Tyrannosaurus </i>FMNH PR2081 ilium and sacrum (after Brochu, 2003). Note even the latter classic theropod has significant medial walls on the peduncles.<br /></td></tr></tbody></table><br /><p>But what does any of this have to do with birds not being dinosaurs? Feduccia spends lots of time discussing the amount of acetabular closure in modern birds, always emphasizing the medially walled areas, and going so far as to say "it is more difficult to adequately assess acetabular morphology in extant birds than is generally realized. Preparatory methods used to produce skeletal collections in museums (e.g., defleshing with dermestid beetles, oxidation methods involving HO, and ammonia, chemical bleach, and maceration) can damage or destroy fragile anatomical features" so that we can't even trust our own specimens! It's as if in his typological mind, this is making birds "less dinosaurian" because dinosaurs are things with open acetabula, and shouldn't this be concerning to us BADists?! No. No it should not. Hell, all crown birds could have fully closed acetabula like derived ankylosaurs and it wouldn't matter one bit because key characters aren't a thing and evolution happens. Just like it doesn't make ankylosaurs any less dinosaurian. We wouldn't even need the fact that every single stem bird fossil* has an open acetabulum regardless of preservation, and that makes it undeniable that regardless of the condition in crown birds, and regardless of what birds evolved from, Mesozoic birds have open acetabula.</p><p>* Even scansoriopterygids which he lies about, claiming "they have closed acetabula" when even his previous description with Czerkas (2014) cited "the partially open acetabulum in <i>Scansoriopteryx</i>." Czerkas and Yuan's (2002) original description also states "the indication from the texture and color extending from the ilium suggests that the hip socket was not as widely perforated as in theropods or dinosaurs in general. The inner edge of this reduced perforation in the hip socket can be seen in both acetabula on the counterslab."<br /></p><p>Feduccia will emphasize "partially closed" in many examples, but non-pennaraptoran theropods are not the monolith he takes them for when as in 2014 he and Czerkas write "theropod dinosaurs, invariably exhibiting a completely perforated and open acetabulum", or as he says in the present paper "The condition in the large Jurassic tetanurine <i>Allosaurus </i>(Figure 3) may be taken as stereotypical of its development in Theropoda", with Figure 3 being redrawn from the 1880s. Just check out classic theropod <i>Ornitholestes </i>above with a medial wall taking up almost half of the acetabular circle. Even in <i>Allosaurus </i>itself, if you look at an actual photo you can see that Figure 3's seemingly uninturrupted dorsal edge is wrongly drawn and that half the ischial peduncle is medial as is some of the pubic peduncle but most of that is hidden by the laterally overlapping supracetabular crest except for the posteroventral corner. Dinosaurs all over the cladogram were changing the amount of medial acetabular ossification and birds were no exception.<br /><br />And really, the takeaway I got from Feduccia's data is that just like most characters, the development of acetabular closure, supracetabular crest extent and antitrochanter size are labile and prone to homoplasy. They are not integral parts of a locomotory system, and similar taxa can manage movement just fine when these variables are changed. That's why you can have <i>Avimimus portentosus</i> and <i>A. nemegtensis</i> with such drastic differences in antitrochanter development, which Feduccia seems baffled by- "it is not clear why a single genus should be polymorphic for this character state, it is difficult to interpret this character conflict." Similarly Feduccia says "It is surprising, given the<br />frequent characterization of birds as having — like dinosaurs — completely open acetabula to<br />discover that rotisserie [chicken] specimens have partially closed acetabula with a perforation covered<br />by soft tissue." Are we assuming Mesozoic dinosaur acetabular foramina were not covered by soft tissue? And is there really a functional difference between the paper thin medial bone walls Feduccia is worried might be present in modern birds but destroyed by preparation and a fibrocartilage wall hypothetically present in a theropod with an open bony acetabulum? Based on Feduccia's described variation within modern birds, I'm doubtful. But he's so obsessed with these being typologically enforced key characters essential to function and evolutionary history, for both himself and his BAD straw man, that them being not all that important in anathema to him.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgLlPFVLbH8P0thQdaJqQdriSFIMJI4f8aMIkwOCuXZqTIOxiz9bOHqOADyS-b8qP2RF3-77fPUX8ewnEV6GrllHVsX4XatulUwvLJCWp3ITUiy9wOqjO5Jm1RCCKNEFUZGn0aEAgnj9CA8UpW488DM0OANph-MQw_sQCAaHnjE9BfAT7Wa2O4nCu2TQ94/s1280/FeducciaFakePelves.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="812" data-original-width="1280" height="203" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgLlPFVLbH8P0thQdaJqQdriSFIMJI4f8aMIkwOCuXZqTIOxiz9bOHqOADyS-b8qP2RF3-77fPUX8ewnEV6GrllHVsX4XatulUwvLJCWp3ITUiy9wOqjO5Jm1RCCKNEFUZGn0aEAgnj9CA8UpW488DM0OANph-MQw_sQCAaHnjE9BfAT7Wa2O4nCu2TQ94/s320/FeducciaFakePelves.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Now, this is out of order but I wanted another figure to liven up the post. On the left is "Late Jurassic <i>Archaeopteryx </i>(image composites of varied sizes, primarily after P. Wellnhofer, based primarily on the Munich specimen, with details from the London, Berlin, and Eichstitt specimens;". Yeah, there is no closing parenthesis and I think "image composites of varied sizes" refers to the <i>Gansus</i>, <i>Qiliania </i>and <i>Pterygornis </i>pictures featured elsewhere in this figure. More importantly as noted far above, this figure is just a drawing (by K. Grow apparently) of a hypothetical composite, and while it is used as an argument against a supracetabular crest in the genus it is incapable of showing the development of a supracetabular crest directed at the viewer as in e.g. <i>Ornitholestes</i>. On the right is an item labeled "<i>Microraptor</i>, the Early Cretaceous four-winged bird-like glider (dromaeosaur), considered by many an early bird. (<i>Microraptor </i>photo courtesy D. Burnham)", but some digging finds this is actually the structure labeled "Right lateral view of microraptoran pelvis (modified cast)" in Burnham's (2007) thesis. The latter claims it is "based on a recent transfer preparation of a Chinese microraptorine, which exposes a lateral view of the acetabulum." Who knows what parts were modified and we have no traceable specimen to confirm anything with, although the preacetabular process and ischial tip don't look like those in actual <i>Microraptor </i>fossils. I'm not even implying any purposeful anatomical inaccuracy, just noting that Feduccia passed off a "modified cast" as an apparently real specimen.<br /></td></tr></tbody></table><br /><p>In the end, Feduccia admits that if "the first birds — and at least taxa like the microraptorines — were not terrestrial cursorial bipeds, like theropods, but trunk-climbing gliders" it "would be consistent with arguments that (at least some of) these taxa are “neoflightless” theropods, whose immediate ancestors underwent transformations in the theropod locomotory system (perhaps because of a shift to arboreality)." But he rejects this, in part due to "the discovery of a “tetrapteryx” stage in early aviation evolution", which is a Petersian thing to say because not even the relatively well resolved pennaraptoran consensus phylogeny views the tetrapteryx stage as an established thing, let alone Feduccia's wacky world of uncertainty where who knows how oviraptorosaurs, troodontids, eudromaeosaurs, microraptorians, unenlagiids, anchiornithines and archaeopterygids interrelate. Sure microraptorians have big leg wings, and <i>Sapeornis </i>and anchiornithines have a much shorter version, but the former are dromaeosaurids and the latter are maybe archaeopterygids, maybe avialans, maybe troodontids. The structures are lacking in <i>Caudipteryx</i>, troodontids, <i>Archaeopteryx </i>and pygostylians, so in order for a tetrapteryx stage to exist, dromaeosaurids and/or anchiornithines would have to be ancestral to crown birds somewhere or we have a lot of reversals. Note Feduccia claims "if <i>Jinfengopteryx </i>is, in fact, a basal troodontid [140,236] (but see discussions in [115,139,162]) then troodontids also primitively exhibited a “tetrapteryx” bauplan", but <i>Jinfengopteryx </i>has no long leg feathers at all, not even long tibiotarsal ones like <i>Archaeopteryx </i>and basal pygostylians. So that's just him being wrong again.</p><p>And in part the rejection is due to... it's really not made clear. Feduccia assumes us BADists are as constrained by imagined evolutionary just-so stories as he is, claiming we are bound to "the argument — advanced since Huxley, championed by Ostrom, repeated ad infinitum — that the obvious reason birds are obligate bipeds and that they never passed through an evolutionary phase in which all four limbs were integrated into the flight mechanism, is that birds are descended from obligate bipeds, viz. theropods." But he already said a facultatively bipedal stage (around the paravian stem maybe?) due to arboreal habits would still be consistent with a theropod ancestry, as quoted in the prior paragraph. Feduccia continues "These data instead suggest that the reason bipedalism in birds differs fundamentally from bipedalism in all other archosaurs [does it?] ... is that birds started their evolutionary trajectory toward obligate bipedalism from a different starting point than the system employed to such success by dinosaurs." But poposaurs and dinosaurs (or alternatively saurischians and <i>Lesothosaurus</i>+genasaurs) started their obligate bipedalism trajectory via facultative bipeds with a more sprawling stance, more closed acetabula, no antitrochanter and less inturned femoral heads just as Feduccia believes birds did. It's just special pleading to claim they are "fundamentally different." Finally, he claims "This is a logical explanation for the surprising persistence of medial occlusion of the acetabulum in many pennaraptorans and the absence, in most, of either well-defined supracetabular crests or antitrochanters." This despite explicitly saying two pages ago "Oviraptorosaurs uniformly display fully perforate acetabula" and "Troodontids have fully perforated acetabula", so by "many pennaraptorans" Feduccia really just means dromaeosaurids. He also said on the previous page "Compsognathids are “prototypical” small theropods [221], yet they lack a supracetabular crest or antitrochanter", so why would this be surprising in most pennaraptorans? (lacking a hypertrophied antitrochanter like ornithurines at least...)</p><p>Amusingly enough, the most vehement objections to BAD are in Feduccia's Figure 7 caption showing hesperornithid pelves. This claims without supporting data "the acetabula show that they are definitively not allied with dinosaurs" and "Hesperornithiforms are often termed diving dinosaurs, but given their acetabula anatomy this designation is erroneous." I'd think such late and specialized divers would be of little relevance, but what do I know... Hey that long and low postacetabular process is looking pretty similar to <i>Postosuchus</i>. Maybe we've found our archosaurian avian ancestor ;)<br /></p><p>And that's it. Feduccia's Conclusion is "The hypothesis that birds are maniraptoran theropod dinosaurs, despite the certitude with which it is proclaimed, continues to suffer from unaddressed difficulties [29-31,50,237]", with references 29-31 being his own books and paper, 50 we recall is James and Pourtless' terrible TERRIBLE attempt at a cladistic analysis, and 237 is <a href="https://www.researchgate.net/publication/351976809_How_Many_Dinosaurs_Are_Birds">James' fawning review of Feduccia's latest book</a>. Ha. You know what's an unaddressed difficulty in Feduccia's hypothesis? Thirty-plus years since he publically doubted the dinosaurian origin of birds, we still have pennaraptorans emerging from some mysterious lineage of Triassic archosaur(-iform)s. And yet we've closed the gaps so well in theropod phylogeny that the precise relationships among basal avialans, basal paravians, basal pennaraptorans, basal maniraptorans/-iforms, basal neocoelurosaurs, basal coelurosaurs, etc. are controversial due to homoplasy. Surely any day now we'll find the evolutionary intermediates between Middle Jurassic scansoriopterygids and Triassic ???????. <b>A</b>nything <b>B</b>ut a <b>S</b>mall <b>R</b>unning <b>D</b>inosaur indeed.<br /></p><p><b>References</b>- Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera <i>Antrodemus</i> (<i>Allosaurus</i>)
and <i>Ceratosaurus</i>. Bulletin of the United States National Museum. 110,
1-154. <br /><br />Harvey, Kaiser and Rosenberg, 1968. An atlas of the domestic turkey (<i>Meleagris gallopavo</i>) myology and osteology. United States Atomic Energyu Commission. 247 pp.<br /><br />Raath, 1977. The anatomy of the Triassic theropod <i>Syntarsus rhodesiensis</i>
(Saurischia: Podokesauridae) and a consideration of its biology. PhD thesis.
Rhodes University. 233 pp.<br /><br />Butendieck, 1980. Die Benennung des Skeletts beim Truthuhn (<i>Meleagris gallopavo</i>) unter Berücksichtigung der Nomina Anatomica Avium (1979). MD vet thesis. Tierärztliche Hochschule Hannover. 153 pp.<br /><br />Chatterjee, 1985. <span style="font-style: italic;">Postosuchus</span>,
a new thecodontian reptile from the Triassic of Texas and the origin of
tyrannosaurs. Philosophical Transactions of the Royal Society of
London, Series B. 309, 395-460.<br /><br />Colbert, 1989. The Triassic dinosaur <i>Coelophysis</i>. Museum of Northern
Arizona Bulletin. 57, 1-174.<br /><br />Baumel and Witmer, 1993. Osteologia. In Baumel, King, Breazile, Evans and Vanden Berge (eds.). Handbook of avian anatomy: Nomina anatomica avium. Second edition. Publications of the Nuttal Ornithological Club. 23, 45-132.<br /><br />Long and Murry, 1995. Late Triassic (Carnian and Norian)
tetrapods from the southwestern United States. New Mexico Museum of Natural History
and Science Bulletin. 4, 1-254.<br /><br />Novas, 1996. Dinosaur monophyly. Journal of Vertebrate Paleontology. 16, 723-741.<br /><br />Benton, 1999. <span style="font-style: italic;">Scleromochlus taylori</span> and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society of London B. 354, 1423-1446.<br /><br />Downs, 2000. <i>Coelophysis bauri</i> and <i>Syntarsus rhodesiensis</i> compared,
with comments on the perparation and preservation of fossils from the Ghost
Ranch <i>Coelophysis</i> Quarry. New Mexico Museum of Natural History and Science
Bulletin. 17, 33-37. <br /><br />Makovicky and Dyke, 2001. Naive falsification and the origin of birds: A commentary. In Gauthier and Gall (eds.). New
Perspectives on the Origin and Evolution of Birds: Proceedings of the
International Symposium in Honor of John H. Ostrom. Yale University. 501-509.<br /><br />Czerkas and Yuan, 2002. An arboreal maniraptoran from northeast China. Feathered
Dinosaurs and the Origin of Flight. The Dinosaur Museum Journal. 1, 63-95. <br /><br />Paul, 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs
and Birds. Johns Hopkins University Press. 460 pp.<br /><br />Sereno, Rao and Li, 2002. <i>Sinornis santensis</i> (Aves: Enantiornithes) from
the Early Cretaceous of northeastern China. In Chiappe and Witmer (eds.). Mesozoic Birds - Above the Heads of Dinosaurs. University of California
Press. 184-208.<br /><br />Weinbaum, 2002. <span class="dont-break-out">Osteology and relationships of <i>Postosuchus kirkpatricki</i> (Archosauria: Crurotarsi). MS Thesis. Texas Tech University. 78 pp.<br /><br /></span>Brochu, 2003. Osteology of <i>Tyrannosaurus rex</i>: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
Society of Vertebrate Paleontology Memior. 7, 138 pp. <br /><br />Burnham, 2007. Paleoenvironment, paleoecology, and evolution of maniraptoran "dinosaurs". PhD thesis. University of Kansas. 183 pp.<br /><span class="dont-break-out"><br />Ferigolo and Langer, 2007 (online 2006). A Late Triassic dinosauriform from south Brazil and the origin of the ornithischian predentary bone. Historical Biology. 19, 23-33.<br /><br /></span>James and Pourtless, 2009. Cladistics and the origin of birds: A review
and two new analyses. Ornithological Monographs. 66, 78 pp.<br /><span class="dont-break-out"><br /></span>Walker and Dyke, 2010 (as 2009). Euenantiornithine birds from the Late Cretaceous of El
Brete (Argentina). Irish Journal of Earth Sciences. 27, 15-62.<br /><span class="dont-break-out"><br /></span>Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae
(Theropoda, Maniraptora)? Anais da Academia Brasileira de Ciencias. 83(1),
117-162. <br /><br />Czerkas and Feduccia, 2014. Jurassic archosaur is a non-dinosaurian bird. Journal
of Ornithology. 155(4), 841-851.<br /><br />Feduccia and Czerkas, 2015. Testing the neoflightless hypothesis: Propatagium
reveals flying ancestry of oviraptorosaurs. Journal of Ornithology. 156(4),
1067-1074.<br /><span class="dont-break-out"><br /></span>Cabreira, Kellner, Dias-da-Silva, da Silva, Bronzati, de Almeida Marsola, Müller, de Souza
Bittencourt, Batista, Raugust and Carrilho, 2016. A
unique Late Triassic dinosauromorph assemblage reveals dinosaur
ancestral anatomy and diet. Current Biology. 26(22), 3090-3095.<br /><br />Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt Basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.<br /><br />Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
<br /><span class="dont-break-out"><br /></span>Müller and Garcia, 2020. A paraphyletic 'Silesauridae' as an
alternative hypothesis for the initial radiation of ornithischian
dinosaurs. Biology Letters. 16(8), 20200417.<br /><br />Feduccia, 2023. The avian acetabulum: Small structure, but rich with illumination and questions. Diversity. 16, 20.<br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com11tag:blogger.com,1999:blog-3248412803814730250.post-33733248388605374872023-10-27T09:22:00.000-07:002023-10-27T09:22:44.797-07:00SVP's big mistake to make a separate virtual conference this year<p><span style="font-size: small;">Due to the pandemic, SVPs 2020 and 2021 were great for people like me who can't afford to travel to it, pay for the registration and lodging, and miss work for that long (I did go in person back in 2001 and 2002 when I unwisely used student loans to afford it... still paying those off...). We got to see all the cool new discoveries for a few hundred dollars and could do so at our leisure without missing concurrent talks, rewatch things we missed, and talk with the presenters without the physical hassles. Then last year they decided to make the online portion optional, so that most of the people with talks I wanted to see didn't upload anything and if I recall correctly, only two of the theropod sessions' presentations were even available. But at least the posters were all there to view, since those require basically no effort to upload unlike narrating a Powerpoint presentation. It was still very disappointing for paying the non-member's United States-based fee of $300.<br /></span></p><p><span style="font-size: small;">This year, SVP decided to make a separate virtual conference from its old-style physical conference. I was hesitant after seeing most of the power players back out of showing anything last year when given the option (I still have no idea what Ruebenstahl et al. proposed about <i>Coelurus</i>...), but you don't know what you're missing if you don't try it. So I paid the slightly lower fee of $200 and yesterday the virtual meeting started and ... what a waste.</span></p><p><span style="font-size: small;"></span></p><p><span style="font-size: small;">The <a href="https://vertpaleo.org/2023-virtual-meeting/">website</a> states "</span>We will still have posters, but they will be in an organized virtual poster hall you can walk around and explore." But what we didn't know until the abstract book came out (conveniently when refunds were also no longer possible) is that only 38 of these are "virtual posters" and that everyone else had the option to let their poster be uploaded but didn't have to. And what we didn't know until yesterday is that only 92 of the 323 normal poster authors agreed to upload their posters to the virtual meeting. That's 28% and includes a whopping FOUR Mesozoic theropod posters, only two of which (the Wessex megaraptoran braincase and the Yellow Cat allosauroid teeth) I really care about. Didn't get Cruz Vega et al.'s Hell Creek avian, Hall et al. on <i>Bistahieversor </i>pneumaticity, Rhynard et al. on <i>Allosaurus jimmadseni</i> internal skull anatomy, Bugos and McDavid on juvenile <i>Coelophysis</i>, Voris et al on <i>Atrociraptor </i>frontals, Takasaki et al. on troodontid intermandibular joints or Perry et al. on <i>Nanuqsaurus </i>validity. Not that those would have justified the $200 price point either (for comparison last year's "very disappointing" virtual meeting still had about fifteen interesting theropod presentations/posters), but it would have been better!</p><p></p><p>Of course the big missing piece was having none of the 305 talks uploaded. No Coppock et al. or Carr and Brusatte on <i>Daspletosaurus </i>species, Slowiak-Morkovina et al. on <i>Bagaraatan</i>, Makovicky et al. on a giant Mussentuchit caenagnathid, Lamanna et al. on unenlagiine <i>Imperobator</i>, Ruebenstahl et al. on <i>Velociraptor </i>species, Maddox et al. on a Hell Creek unenlagiine or Hohman et al. on Two Medicine dromaeosaurid variation, and that's just page 56 of the abstract volume. Not only were most of the most interesting presentations given as talks, but a talk is far more engaging than a poster. It's ten to fifteen minutes long, whereas even the most text-dense posters take maybe a couple minutes to read, and most much less. Similarly, talks are more entertaining than posters, which means I ended up watching a lot of talks from SVPs 2020-2022 that were outside my area of expertise, and learned more. But I have about as much interest in reading a poster on unfamiliar material as I do reading a paper on it. So instead of a potential ~61 hours (305 talks times 12 minutes) of entertainment just from talks alone, we got 130 posters which took a few hours to get through. <b>For comparison, that was ~70 hours for $300 in 2020 and 2021 versus ~3 hours for $200 this year.</b></p><p>Finally, SVP got the Q&A part down perfectly in 2020 and 2021 with a chatroom for each presentation. In real life, you have to either awkwardly raise your hand for the limited minute or so after the talk and hope to be called on or track down the author during the meeting and hope they're not having a more important conversation if you find them. But in 2020 and 2021, if you have any question you can just post it and the author will see it at some point and reply, with time to formulate a good answer. Now in 2023 we have the virtual Sternberg Museum where we have to talk in real time while our avatars are next to each other and any author may be there or may not be at any given time, and if they aren't one of the authors of the 38 virtual posters have no obligation to be there at all. And just like in real life, if you do happen to track one down they may be in one of the numerous virtual tables that are private chat rooms, so I guess you watch your avatar wait for them to get up? Or maybe it's possible to interrupt the private chat awkwardly? After checking, not even the virtual poster authors have any obligation, as they are merely "welcome to stand with their poster at any time". With the average abstract having three authors are we supposed to keep track of all the authors of all the abstracts we are interested in and keep checking the Participants bar to potentially chat with them? It'd be simpler just to email or Facebook message authors at this rate. It reminds me of <a href="https://www.youtube.com/watch?v=u6UYGwPaLf8">that meme of shopping in the metaverse</a> where you have to push a virtual cart and turn to pick up virtual items instead of just selecting from a list like every online retailer - SVP has recreated the bad parts of in person Q&A for its virtual world.</p><p>"But wait!", you say. What about the online components that aren't talks or posters? They exist in hour or two hour intervals, and weren't announced until 10 pm Wednesday, approximately zero days before the meeting began. So while it would be easier to find participating dinosaur authors at the '<span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">Networking Social: Dinosaur Research' between 6 and 8 pm Friday, I gotta work and had no time to plan around it. Besides that, '</span><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">Discussion Panel: State of the Dinosauria' might be interesting to watch (happening the hour before I work Friday, so I'll get to be there for the first 45 minutes), but this and the other Zoom presentations were stated to be uploaded to YouTube later so aren't really perks of attending the meeting.</span></p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiBFbE2uNufA0aMwbseOnP21OyRhMmU6yausqXeoDBEPclppYkc1uVdYjC6l8PvjDv42mNUAlx5n6AoXGOpE17LFRCOFoHX7MKcbQL7h3HUVRem3PwoqhrC6WGX93shpssbO7mUQzP0aiaCUlcBzhyJgslnxPqTe3Kp3ieaHoLLhxgEZK6KDGXqL6kYIN4/s3430/SVP2023.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1306" data-original-width="3430" height="122" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiBFbE2uNufA0aMwbseOnP21OyRhMmU6yausqXeoDBEPclppYkc1uVdYjC6l8PvjDv42mNUAlx5n6AoXGOpE17LFRCOFoHX7MKcbQL7h3HUVRem3PwoqhrC6WGX93shpssbO7mUQzP0aiaCUlcBzhyJgslnxPqTe3Kp3ieaHoLLhxgEZK6KDGXqL6kYIN4/s320/SVP2023.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Waiting for Choiniere's talk 30 minutes before it began, but I don't think it was even here and I had to use a link from an email instead.<br /></td></tr></tbody></table><br /></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">Thus while <a href="https://vertpaleo.org/2023-registration-information/">$740</a> gets you up to 25 hours of talks (since in real life there are three sets of twenty-six 15 minute talks happening simultaneously every day) and 453 posters (let's say at 2 minutes per poster that's 9 hours to digest them all) plus all the in person events and online events, $200 gives you 130 posters that might take three hours to digest and the same online events. <b>Even ignoring the in person events and quality of talks over posters, you're getting 9% of the time from the online meeting while spending 27% of the cost. $740 times 9% is $67. A pretty far cry from $200.</b></span></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">Honestly, for the last three SVPs I've spent days viewing everything I wanted to whereas this year I was done in a few hours of virtual walking to each poster in order. I watched Choiniere's Elliot Formation talk right before posting this (which was fine), will catch most of the dinosaur panel before work Friday, then probably watch the squamate panel Saturday and mammal panel Sunday because I want to feel some sort of relevance for my expenditure, but that's literally everything I'm remotely interested in. I'd say the value is <u>maybe </u>$30. It's sad.</span></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;"><br /></span></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">Let's end this on a positive note though - how to improve things for future years. The number one message that should have obviously been learned from this year and last is to UPLOAD EVERY TALK AND POSTER. I get that not everyone has the equipment or skills, so just have SVP record the live talks and post those with an option for the authors to upload a high resolution version of the slideshow too. To make it more engaging, have the presentation sessions from each room be streamed live via Zoom, then the talks broken up to be watched separately later. For those 72% of presenters who are apparently paranoid someone is going to somehow get away with stealing your ever so tweet-worthy study on <i>Deinonychus </i>histology or whatever, I'm pretty sure nothing has been claim-jacked from SVPs 2020 or 2021 despite people having years to do so. What's ironic is that SVP solved one of its main problems with technology- there are too many talks to view them all in person because they have to host three sets simultaneously, but by hosting them all online as videos on demand everyone can watch all of them. Then this year they killed that solution and made the world worse for all the attendees with a return to artificial scarcity. The second message is bring back the chatroom Q&A. It was perfect and now you've recreated all the bad parts of in person interactions with new negatives like authors not even needing to be in the available space. So again you've solved the real life problems with technology then just brought them back for no reason.</span></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;"><br /></span></p><p><span class="OYPEnA" style="color: black; font-style: normal; font-weight: 400;">COVID 19 forced SVP to update to the Shiny Digital Future, but now it's backsliding to be more tedious for everybody. Even the SVP members who get paid to attend via their institutions now again have to decide what talk room to attend, can't watch talks on their own time, can't rewatch talks, have to track down authors, etc.. 2020 and 2021 showed us you can deliver a quality digital conference, please do it again in 2024. Or if you insist on being as archaic as your material of interest, charge an order of magnitude less for that option.<br /></span></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com0tag:blogger.com,1999:blog-3248412803814730250.post-47644675424108333302023-07-09T16:21:00.000-07:002023-07-09T16:21:50.994-07:00New data on African abelisaurs<p>Hi everyone. I'm a coauthor on a paper published this week reviewing the record of African abelisauroids (de Souza-Júnior et al., 2023). I want to thank André Luis de Souza-Júnior for adding me, as he used The Theropod Database as a resource. That's how you do it! We incorporated my corrections and suggestions back in 2020 before it was submitted, but as is typical of these projects there have been new discoveries since. Here are some additional records-<br /><br /></p><p><b>unnamed Abelisauroidea</b> (Lasseron, 2020)<br />
<b>E</b><span style="font-weight: bold;">arly Bathonian, Middle Jurassic</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">GEA 6, Guelb el Ahmar,
Anoual Formation, Morocco<br />
Material</span>- (MNHN GEA6-5) lateral tooth (2.58x1.86x1.39 mm)<br />
<b>E</b><span style="font-weight: bold;">arly Bathonian, Middle Jurassic</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">GEA 7, Guelb el Ahmar,
Anoual Formation, Morocco<br />
</span>(MNHN GEA7-14) lateral tooth (3.75x2.51x1.59 mm)<br />
(MNHN GEA7-17) lateral tooth (2.60x2.06x1.09 mm)<br />
<b>Comments</b>- Discovered in 2015 and/or 2018, Lasseron (2020) concluded "<span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">The convexity of the two carinae is a remarkable feature.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">It
recalls the Abelisauridae (Smith, 2007; Hendrickx & Mateus, 2014),
but also the Noasauridae (Carrano et al., 2002), an identification
which would be consistent with the small size of these teeth.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">We therefore attribute these teeth to an Abelisauroidea gen.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">and sp.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">indet." (translated).</span></span></span><br />
<b>Reference</b>- Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.<br />
</p>
<p><span style="font-weight: bold;">unnamed abelisauroid</span> (Lasseron, 2020)<br />
<b>Berriasian, Early Cretaceous<br />KM-A1, </b><b>Ksar Metlili, </b><b>Ksar Metlili Formation, Morocco<br />
Material</b>- (FSAC-KM-A1-12) lateral tooth (3.41x2.26x1.47 mm)<br />
<span style="font-weight: bold;">Comments</span>- Collected in 2010, 2015 or 2018, Lasseron (2020) concluded "<span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">The convexity of the two carinae is a remarkable feature.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">It
recalls the Abelisauridae (Smith, 2007; Hendrickx & Mateus, 2014),
but also the Noasauridae (Carrano et al., 2002), an identification
which would be consistent with the small size of these teeth.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">We therefore attribute these teeth to an Abelisauroidea gen.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">and sp.</span></span> <span class="jCAhz ChMk0b"><span class="ryNqvb">indet." (translated).<br />
<span style="font-weight: bold;">Reference</span>- </span></span></span>Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.<br />
<br />
<b>undescribed possible abelisauroid</b> (Smith, Lamanna, Dodson, Attia and Lacovara,
2001)<br />
<b>Cenomanian, Late Cretaceous<br />Bahariya Oasis, Baharija Formation, Egypt</b><br />
<b>Material</b>- teeth (FABL 7.5 mm)<br />
<span style="font-weight: bold;">Comments</span>- These "recently recovered" teeth are stated to have significantly more serrations per mm than similarly sized <span style="font-style: italic;">Deinonychus</span> or <span style="font-style: italic;">Dromaeosaurus</span>
and have "denticle morphologies and the average crown base width (CBW)
and CBL9crown base length] relationships ... more reminiscant of
abelisaurids than dromaeosaurids."<br />
<b>Reference</b>- Smith, Lamanna, Dodson, Attia and Lacovara, 2001. Evidence
of a new theropod from the Late Cretaceous of Egypt. Journal of Vertebrate Paleontology.
21(3), 102A.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9O2venTxYXK5gO_h7O6H4zdqeAhIzPEusTKwwlzu7gxNsz0fJt1nmqltdCDd1eJDlvSxpg1FKfCByI8kDeKu06uWiIsF6ZJeN3_8zFYie1067OXUsyXeRjb0Fo3uIvb8GEShZrLvb4XynwFWlrJIFV86F-H41RHi8NXZamrEQVGWFrjOEyoSJyJNCCSM/s1823/Kem%20Kem%20noasaurid%20cervical.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1257" data-original-width="1823" height="221" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9O2venTxYXK5gO_h7O6H4zdqeAhIzPEusTKwwlzu7gxNsz0fJt1nmqltdCDd1eJDlvSxpg1FKfCByI8kDeKu06uWiIsF6ZJeN3_8zFYie1067OXUsyXeRjb0Fo3uIvb8GEShZrLvb4XynwFWlrJIFV86F-H41RHi8NXZamrEQVGWFrjOEyoSJyJNCCSM/s320/Kem%20Kem%20noasaurid%20cervical.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Kem Kem noasaurid cervical FSAC-KK-5016 in (A) right lateral, (B) left lateral, (C) anterior, (D) dorsal, (E) ventral and (F) posterior views (after Smyth et al., 2020). Scale = 10 mm.<br /></td></tr></tbody></table><br /><p><b>unnamed Noasauridae</b> (Russell, 1996)<br />
<b>Cenomanian, Late Cretaceous<br />
Kem Kem beds, Morocco</b><br />
<b>Material</b>- (CMN 41873; bone taxon H) distal humerus (48 mm wide) (Russell,
1996) <br />
(FSAC-KK-5016) (~1.5 m) incomplete ~fourth cervical vertebra (21 mm) (Smyth, Ibrahim, Kao and Martill, 2020)<br />
<b>Comments</b>- The humerus is very similar to <i>Masiakasaurus</i>.<br />
<b>References</b>- Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous
of the Tafilalt, Morocco. Bulletin du Museum national d'Histoire naturelle.
18, 349-402.<br />
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.<br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjkDSoS5ZnEaj31sfRYatSoJve8pm4RwaZaNs1ty7dNCZHNQmLGZXO8agYsmEW_kqFrQBGvg0lY5J0Ng0kMfmkwFFoVTCuh5E7LCXXOG6J_4DEaF0n_Q7L1RPkaaMmnPkAFO_SFbfftnr_3AHXh7711JOEsCiNcBxD0RhKktIWqnraUdJ437G9LPu_w8pI/s1824/Kem%20Kem%20abelisaurid%20axis.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1174" data-original-width="1824" height="206" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjkDSoS5ZnEaj31sfRYatSoJve8pm4RwaZaNs1ty7dNCZHNQmLGZXO8agYsmEW_kqFrQBGvg0lY5J0Ng0kMfmkwFFoVTCuh5E7LCXXOG6J_4DEaF0n_Q7L1RPkaaMmnPkAFO_SFbfftnr_3AHXh7711JOEsCiNcBxD0RhKktIWqnraUdJ437G9LPu_w8pI/s320/Kem%20Kem%20abelisaurid%20axis.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Kem Kem abelisaurid axis FSAC-KK-5015 in (A) right lateral, (B) left lateral, (C) anterior, (D) dorsal, (E)
ventral and (F) posterior views (after Smyth et al., 2020). Pink area is filler. Scale = 20 mm.<br /></td></tr></tbody></table><br /><p><b>unnamed Abelisauridae</b> (Fanti, Cau, Martinelli and Contessi, 2014)<br />
<b>Cenomanian, Late Cretaceous<br />
Kem Kem beds, Morocco<br />
Material</b>- (CMN 50446) tooth (~24x~11x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />?(FSAC-KK 912) tooth (~17x~10x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
?(FSAC-KK 913) tooth (~22x~12x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
?(FSAC-KK 914) partial tooth (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
?(FSAC-KK 915) tooth (~18x~8x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
(FSAC-KK-5015) (~2.7 m) axis (38 mm) (Smyth, Ibrahim, Kao and Martill, 2020)<br />?(FSAC-KK unnumbered) tooth (~16x~11x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
(MNHN MRS 348) tooth (48x17x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)<br />
(MNHN MRS 783) tooth (28x14x6 mm) (Fanti, Cau, Martinelli and Contessi, 2014)<br />
(MNHN MRS 1266) tooth (34x14x7 mm) (Fanti, Cau, Martinelli and Contessi, 2014)<br />
(MNHN MRS 1838) tooth (36x19x9 mm) (Fanti, Cau, Martinelli and Contessi, 2014)<br />
(MSNM V6053) tooth (~23x~10x? mm) (Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020)<br />
<span style="font-weight: bold;">Comments</span>- Fanti et al. (2014) list
measurements of four "Abelisauridae indet." teeth housed in the MNHN
which have MRS numbers corresponding to the Gara Sbaa locality (e.g. <span style="font-style: italic;">Rebbachisaurus</span> type) of the Kem Kem beds. Smyth et al. suggested the axis
FSAC-KK-5015 represents an abelisaurid outside
Majungasaurinae+Brachyrostra, perhaps a juvenile <span style="font-style: italic;">Rugops</span>. Ibrahim et al. (2020) figure CMN 50446 and MSNM V6053 as "?Abelisaurid
tooth", and several others as "Indeterminate theropod teeth" which seem
to be abelisaurid based on the low distal curvature in each, and
marginal undulations in FSAC-KK 912 and 915.<br />
<b>References</b>- Fanti, Cau, Martinelli and Contessi, 2014. Integrating palaeoecology
and morphology in theropod diversity estimation: A case from the Aptian-Albian
of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology. 410, 39-57.<br />
Ibrahim, Sereno, Varricchio, Martill, Dutheil, Unwin, Baidder, Larsson,
Zouhri and Kaoukaya, 2020. Geology and paleontology of the Upper
Cretaceous Kem Kem Group of eastern Morocco. ZooKeys. 928, 1-216.<br />
Smyth, Ibrahim, Kao and Martill, 2020 (online 2019). Abelisauroid
cervical vertebrae from the Cretaceous Kem Kem beds of southern Morocco
and a review of Kem Kem abelisauroids. Cretaceous Research. 108, 104330.<br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi_8P8twKLGxbM9V1V_XUc2MAeLecwFhtaZD_rzTCg60wt9dSZdNzTLFSF7bVnRDnIxsGfxwgiXugTjiVDZqnTkhKNkVlq7uSy8vCvuulH3U2X-4MXDlPEVm0tG2KCtbRG2f-7IgAYGOmpCqUGBz7zRTjRlpWoDyPiidr1OQ-tkFqkbEIBFvukAh4pWjS8/s1625/Quseir%20abelisaurid%20fibula.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="956" data-original-width="1625" height="188" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi_8P8twKLGxbM9V1V_XUc2MAeLecwFhtaZD_rzTCg60wt9dSZdNzTLFSF7bVnRDnIxsGfxwgiXugTjiVDZqnTkhKNkVlq7uSy8vCvuulH3U2X-4MXDlPEVm0tG2KCtbRG2f-7IgAYGOmpCqUGBz7zRTjRlpWoDyPiidr1OQ-tkFqkbEIBFvukAh4pWjS8/s320/Quseir%20abelisaurid%20fibula.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Quseir abelisaurid left fibula in (F) lateral, (G, H) medial and (I, J) proximal views (after Salem et al., 2021). Scale = 20 mm.<br /></td></tr></tbody></table><br /><p><span style="font-weight: bold;">unnamed probable abelisaurid</span> (Sallam, O'Connor, Kora, Sertich, Seiffert, Faris, Ouda, El-Dawoudi, Saber and El-Sayed, 2016)<br />
<span style="font-weight: bold;">Middle Campanian, Late Cretaceous</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">Baris, El Hindaw Member, Quseir Formation, Kharga Oasis, Egypt</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">Material</span>- (MUVP 187) proximal fibula (25x5 mm prox)<br />
<span style="font-weight: bold;">Comments</span>- Discovered in 2008 or
2010, Sallam et al. (2016) initially reported "A proximal left fibula
of a theropod dinosaur was also collected from the Baris area" and
listed it as Theropoda indet.. Salem et al. (2021) noted the
prominent m. iliofibularis tubercle is like abelisaurids and it has a
"wide, slightly deep [proximomedial] fossa that closely resembles the
condition in the abelisaurid <span style="font-style: italic;">Majungasaurus</span>." They say it "exhibits some affinities with abelisaurids" and refer to it as "Cf. Abelisaurid" in theit table 3. <br />
<span style="font-weight: bold;">References</span>- Sallam, O'Connor,
Kora, Sertich, Seiffert, Faris, Ouda, El-Dawoudi, Saber and El-Sayed,
2016. Vertebrate paleontological exploration of the Upper Cretaceous
succession in the Dakhla and Kharga Oases, Western Desert, Egypt.
Journal of African Earth Sciences. 117, 223-234.<br />
Salem, O'Connor, Gorscak, El-Sayed, Sertich, Seiffert and Sallam, 2021.
Dinosaur remains from the Upper Cretaceous (Campanian) of the Western
Desert, Egypt. Cretaceous Research. 123, 104783.<br /><br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgz-JgnnbGrPSvEkXI5oOiYKHtc6hAgPGlYP8fm9utTl3h0ROjPYlVQ_GtX6qH1E96G7MhjAGkGrqblJeASXTR44I4eS7c8VkEQuvwH9TWYn6VvixC-6SCP6bZC4diXywfxmnFqr4_Z_o5xIrswxDs_s851G7s_RMfHBXrQDmqO2pwm4AJhIdEMm48eYU0/s953/Elrhaz%20abelisaurid%20carina.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="940" data-original-width="953" height="316" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgz-JgnnbGrPSvEkXI5oOiYKHtc6hAgPGlYP8fm9utTl3h0ROjPYlVQ_GtX6qH1E96G7MhjAGkGrqblJeASXTR44I4eS7c8VkEQuvwH9TWYn6VvixC-6SCP6bZC4diXywfxmnFqr4_Z_o5xIrswxDs_s851G7s_RMfHBXrQDmqO2pwm4AJhIdEMm48eYU0/s320/Elrhaz%20abelisaurid%20carina.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Elrhaz abelisaurid carina MNHN F.GDF-M30 in (E) labiolingual and (F) mesiodistal views (after Pochat-Cottilloux et al., 2022). Scale = 1 mm.<br /></td></tr></tbody></table><br /><p><span style="font-weight: bold;">unnamed Abelisauridae</span> (Pochat-Cottilloux, Allain and Lasseron, 2022)<br />
<span style="font-weight: bold;">Aptian, Early Cretaceous</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">GAD 5, Gadoufaoua, Elrhaz Formation, Niger</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">Material</span>- (MNHN F.GDF-M30) two tooth fragments<br />
(MNHN F.GDF-M37) two partial teeth<br />
<span style="font-weight: bold;">Comments</span>- Collected in 1970,
Pochat-Cottilloux et al. (2022) noted "some of this material was stored
at the MNHN in a separate box" and had a "lighter color and a better
quality of fossil preservation than other seen in other samples" which
"suggested a different depositional environment to that of other
samples so, as a result, this sample was separated from the rest" ...
"as "site GADb"." This is true for MNHN F.GDF-M30. The
authors state there are 3 serrations per mm on MNHN F.GDF-M30 and 4 per
mm on MNHN F.GDF-M37, in the latter the carinae "extend along the
entire length of both sides of the base" and in all specimens "the
shape of the denticles and their number are typical of theropod teeth
and could be assignable to Abelisauridae."<br />
<span style="font-weight: bold;">Reference</span>- Pochat-Cottilloux,
Allain and Lasseron, 2022. Microvertebrate fauna from Gadoufaoua
(Niger, Aptian, Early Cretaceous). Comptes Rendus Palevol. 21(41),
901-926.<br /><br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgrI1eoaef7P6OskWa94gJL5wqkhe92QlLzkc-R7spi1sWtYy2h4h_Sr4sIdsUGOG4XtVxOa03YwxduyRGLr_bys86xbZ0x3-EzBy6tSdYDM1ES3E14We4fjtieXReUCi92cpWq1DGVj2GySDpOnw7RE0ssrh9vyS2UTYQRnTa8XuSDhgO3Cf4HC83agNA/s1065/Galula%20abelisaurid%20tooth.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="687" data-original-width="1065" height="206" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgrI1eoaef7P6OskWa94gJL5wqkhe92QlLzkc-R7spi1sWtYy2h4h_Sr4sIdsUGOG4XtVxOa03YwxduyRGLr_bys86xbZ0x3-EzBy6tSdYDM1ES3E14We4fjtieXReUCi92cpWq1DGVj2GySDpOnw7RE0ssrh9vyS2UTYQRnTa8XuSDhgO3Cf4HC83agNA/s320/Galula%20abelisaurid%20tooth.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Galula abelisaurid tooth TNM 02088 in labial view (after O'Connor et al., 2006). Scale = 5 mm.<br /></td></tr></tbody></table><br /><p><b>unnamed Abelisauridae</b> (O'Connor, Gottfried, Stevens, Roberts, Ngasala, Kapilima and
Chami, 2006)<br />
<b>Albian, Early Cretaceous<br />TZ-07, Namba Member of the Galula Formation, Tanzania</b><br />
<b>Material</b>- (TNM 02088) lateral tooth (~19x~11.5x? mm)<br />
(TNM coll.) eight lateral teeth, two anterior teeth<br />
<span style="font-weight: bold;">Comments</span>- Discovered between
2002 and 2005, this material was described as theropod by O'Connor et
al. (2006) and referred to the Unit I of the Red Sandstone Group at the
time. Roberts et al. (2010) subsequently revised the stratigraphic
nomenclature, naming Unit I the Galula Formation. Salem et al. (2021)
refers the isolated teeth to Abelisauridae without comment, which seems
likely given their stratigraphy, locality, size and morphology
(particularly the nearly straight distal edge). Several of the teeth
were associated with the holotype of the titanosaur <span style="font-style: italic;">Shingopana</span>,
"suggesting a possible scavenging event." Note the theropod
caudals from the same locality described by O'Connor et al. differ from
abelisaurids in their strong infradiapophyseal laminae and are here
tentatively referred to Megaraptora.<br />
Teeth have ~13-16 serrations per 5 mm mesially and distally. O'Connor
et al. further indicate "Two teeth are D-shaped in cross-section,
indicating a position in the rostralmost portion of either the dentary
or premaxilla."<br />
<b>References</b>- O'Connor, Gottfried, Stevens, Roberts, Ngasala, Kapilima and
Chami, 2006. A new vertebrate fauna from the Cretaceous Red Sandstone Group,
Rukwa Rift Basin, southwestern Tanzania. Journal of African Earth Sciences.
44, 277-288.<br />
Roberts, O'Connor, Stevens, Gottfried, Jinnah, Ngasala, Choh and
Armstrong, 2010 (online 2009). Sedimentology and depositional
environments of the Red Sandstone Group, Rukwa Rift Basin, southwestern
Tanzania: New insight into Cretaceous and Paleogene terrestrial
ecosystems and tectonics in sub-equatorial Africa. Journal of African
Earth Sciences. 57, 179-212.<br />
Salem, O'Connor, Gorscak, El-Sayed, Sertich, Seiffert and Sallam, 2021.
Dinosaur remains from the Upper Cretaceous (Campanian) of the Western
Desert, Egypt. Cretaceous Research. 123, 104783.<br /><br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgSldVeNgE6xw1ulQYMQ-IWEU6vQnIdGRMO-LCbcjoHq9JKIPyYJmkswPCrAFds6l616TFBd1O9cQuPMxZiXRfEVzf_8oyBzk9yqYGa-ZZwGXa_C2lGDz_O9f1WO0zSIJ5adO9qjMiHolUUzViynkklxCRDsUbaz8cTBTkQTrCUCZRxre1OTDU5QrI_fQ4/s1372/Gokwe%20theropod%20teeth.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="918" data-original-width="1372" height="214" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgSldVeNgE6xw1ulQYMQ-IWEU6vQnIdGRMO-LCbcjoHq9JKIPyYJmkswPCrAFds6l616TFBd1O9cQuPMxZiXRfEVzf_8oyBzk9yqYGa-ZZwGXa_C2lGDz_O9f1WO0zSIJ5adO9qjMiHolUUzViynkklxCRDsUbaz8cTBTkQTrCUCZRxre1OTDU5QrI_fQ4/s320/Gokwe%20theropod%20teeth.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Gokwe abelisaurid (left, right) and averostran (center) teeth (University of Zimbabwe coll.) (after Bond and Bromley, 1970).<br /></td></tr></tbody></table><br /><p><b>undescribed Abelisauridae</b> (Bond and Bromley, 1970)<br />
<b>Early Cretaceous<br />
Gokwe Formation, Zimbabwe<br />
Material</b>- (University of Zimbabwe coll.) at least two teeth (~48x~31x?, ~49x~34x? mm)<br />
<span style="font-weight: bold;">Comments</span>- Bond and Bromley
(1970) were the first to figure three theropod teeth, discovered
between 1962
and 1970, as their Plate 1A "Reptilian teeth of two types from the
Gokwe Formation." Listed as being " in collection of Geology
Department, University College of Rhodesia", the latter has since been
renamed the University of Zimbabwe. The first and third are
clearly abelisaurid, with broad bases, straight to convex distal edges
and fine serrations both mesially and distally (~6 per 5 mm on both
carinae). The second is narrower and more recurved, so referred
to Averostra indet. here.<br />
<b>Reference</b>-
Bond and Bromley, 1970. Sediments with the remains of dinosaurs near
Gokwe, Rhodesia. Palaeogeography, Palaeoclimatology, Palaeoecology.
8(4), 313-327.<br /><br /></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjqR0bDA96AoVyZdDyxPsldUFfjGPkRBltbCQqAQyb5E10wK1CdmVtZ_Hd-Z7NSKGB8hSXsNmmnPaGhNNU2fcSnVOHeZxdy5rc8ruH7b0wtBIA5t__7DmB2WG43-9qGL02Byo67WcT2cwffBAkK2ZXT6K9nhfhWxO_4CmjabLDZ9sp1ZpDr0U7uTC_j09s/s1074/Baharija%20abelisaurid%20cervical.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1074" data-original-width="865" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjqR0bDA96AoVyZdDyxPsldUFfjGPkRBltbCQqAQyb5E10wK1CdmVtZ_Hd-Z7NSKGB8hSXsNmmnPaGhNNU2fcSnVOHeZxdy5rc8ruH7b0wtBIA5t__7DmB2WG43-9qGL02Byo67WcT2cwffBAkK2ZXT6K9nhfhWxO_4CmjabLDZ9sp1ZpDr0U7uTC_j09s/s320/Baharija%20abelisaurid%20cervical.png" width="258" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Baharija abelisaurid tenth cervical vertebra MUVP 477 in (a) anterior, (b) posterior, (c) left lateral, (d) right lateral), (e) ventral and (f) dorsal views (after Salem et al., 2021).<br /></td></tr></tbody></table><p><br /><span style="font-weight: bold;">unnamed possible brachyrostran</span> (Salem, Lamanna, O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2021)<br />
<b>Early Cenomanian, Late Cretaceous<br />Gebel el Dist, Baharija Formation, Egypt<br />
Material</b>- (MUVP 477) (~5.8 m) tenth cervical vertebra (67.00 mm)<br />
<span style="font-weight: bold;">Comments</span>- Discovered in 2016,
Salem et al. (2021) added this to Tortosa et al.'s ceratosaur matrix
and recovered it as an abelisaurid based on "(1) long axis of
diapophysis forms angle of 65° to midsagittal plane; (2) dorsal surface
of neural arch clearly delimited from lateral surface of diapophysis;
(3) deep spinoprezygapophyseal and spinopostzygapophyseal fossae; and
(4) well-developed epipophyses, comparable to those observed in the
tenth cervicals of <span style="font-style: italic;">Majungasaurus</span> and MPM 99 (but smaller than those of <span style="font-style: italic;">Carnotaurus</span> and <span style="font-style: italic;">Ekrixinatosaurus novasi</span>)."
The resulting Salem et al. (2022) publication officially describing the
specimen did not use character 1. While it was a brachyrostran
(in a polytomy with <span style="font-style: italic;">Ekrixinatosaurus</span>, <span style="font-style: italic;">Ilokelesia</span>
and the derived clade) in majority rule trees, a posteriori pruning of
taxa was not done to determine if this is real signal or merely an
artifact. Examination of the 2022 supplementary data should resolve this.<br />
<span style="font-weight: bold;">References</span>- Salem, Lamanna,
O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2021. First
definitive record of Abelisauridae from the Bahariya Formation,
Bahariya Oasis, Western Desert of Egypt increases diversity of
large-bodied theropods in the MIddle Cretaceous of northeastern Africa.
The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 225-226.<br />
Salem, Lamanna,
O'Connor, El-Qot, Shaker, Thabet, El-Sayed and Sallam, 2022. First
definitive record of Abelisauridae (Theropoda: Ceratosauria) from the
Cretaceous Bahariya Formation, Bahariya Oasis, Western Desert of Egypt.
Royal Society Open Science. 9: 220106.<br /><br /><br /><br />Additionally, a review paper isn't really the place for new information or unpublished sources, so here are some extensive updates to poorly documented finds-<br /><br /><b><a name="Sidormimus"></a>"Sidormimus"</b> Molina-Perez and Larramendi, 2019<br />
= "Dogosaurus" Anonymous, 2000 online<br />
<b>Aptian-Albian, Early Cretaceous<br />Gadoufaoua, Elrhaz Formation, Niger</b><br />
<b>Material</b>- (MNN GAD coll.) (~1 m) partial skeleton including maxilla, cervical ribs, dorsal vertebrae,
dorsal ribs, uncinate processes, sacral vertebrae, scapula, coracoid, sternal
plates, sternal ribs, humerus, radius, ulna, carpus, manus including manual
digit II, phalanges, manual unguals and manual claw sheath, pelvis including
pubis and hindlimb including femur, tibia, fibula, calcaneum, metatarsus, pedal digit II, pedal
digit III, pedal digit IV and pedal unguals<br />
<b>Comments</b>-
This specimen was discovered on September 13 2000 and announced by Lyon (2000 online) as "a brand
new, dog-sized theropod, which the team
has affectionately (but unofficially) named
"<i>Sidormimus</i>."" and a "<span class="updateCaption">new,
dog-sized carnivore - perhaps one of the smallest dinosaurs ever", with
a photo of the specimen in situ. </span>The same photo was labeled <span style="font-style: italic;">Dogosaurus</span> on a dispatch from Project
Exploration on the National Geographic website, which also stated it was "<span class="bodycontent">no more than three feet long. Its slender neck
and ribcage were preserved poking out of the rock. Its long, clawed
hind leg was uncovered as we dug around the exposed bones</span>."
Sereno et al. (2004) first announced the
specimen in print, as "a small (1 m) articulated skeleton showing many
abelisauroid and noasaurid synapomorphies, including pneumatized
presacral and sacral neural arches, proportionately long presacral
centra, and others" and "preserves a maxilla and pelvic girdle, the
former of which bears the distinctive abelisaurid pit-and-groove
texturing of the skull bones." They include it in their cladogram
as "Gadoufaoua noasaurid", but it is not an OTU in the matrix. It
is also listed
in Sereno and Brusatte (2008) as "undescribed noasaurid" in a faunal
list, and is noted to have a pubic boot more with limited expansion
than the "<span style="font-style: italic;">Kryptops</span>" postcrania MNN GAD1-2. Sidor
(pers. comm. 2005) confirms the "Sidormimus" specimen is the Elrhaz
noasaurid. Sereno (2010) devoted an SVP abstract to it, noting the specimen has- long and
robust posterior cervical ribs, dorsal centra more than twice as long
as tall, five uncinate processes (unique among non-maniraptoran
theropods), posteriorly directed glenoid, enlarged coracoid,
ossified sternal plates and ribs, forelimb 18% of hindlimb length,
robust deltopectoral crest and olecranon process, manual digit II
longer than III, straight manual unguals, tibia longer than femur,
pedal digits II and IV much shorter than III, and short flat pedal
unguals. Sereno (2017) provided further information- straight
tibia, weak iliofibularis tubercle on fibula, fibula distally fused
with calcaneum, no dorsal vascular groove on pedal unguals. The
co-occuring <span style="font-style: italic;">Afromimus</span>
shared fibular-calcaneum fusion but differs in the other characters, in
addition to having a more robust proximal fibula and distal fibula
which is expanded transversely. Molina-Perez and Larramendi
(2019) first used the name in print, specified to be a nomen nudum.<br />
<b>References</b>- Lyon, online 2000. <a href="http://web.archive.org/web/20121024124915/http://www.projectexploration.org/niger2000/9_15_2000.htm">http://web.archive.org/web/20121024124915/http://www.projectexploration.org/niger2000/9_15_2000.htm</a><br />Anonymous, online 2000.
<a href="http://web.archive.org/web/20001208070300/http://www.nationalgeographic.com/dinoquest/profile_01_dispatch2b.html">http://web.archive.org/web/20001208070300/http://www.nationalgeographic.com/dinoquest/profile_01_dispatch2b.html</a>
<br />
Sereno, Wilson and Conrad, 2004. New dinosaurs link southern landmasses in the
Mid-Cretaceous. Proceedings of the Royal Society, Series B. 271, 1325-1330.<br />
Sereno and Brusatte, 2008. Basal abelisaurid and carcharodontosaurid theropods
from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica.
53(1), 15-46.<br />
Keillor, Sereno and Masek, 2010. Range of movement in a noasaurid forelimb:
In situ data and joint reconstruction. Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 114A.<br />
Sereno, 2010. Noasaurid (Theropoda: Abelisauroidea) skeleton from Africa shows
derived skeletal proportions and function. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 162A.<br />
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.<br />
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.</p><p><b><a name="Titanovenatorkenyaensis"></a>"Titanovenator" </b>Boyd, 2020 online<br />
<span style="font-weight: bold;">"T. kenyaensis"</span> Boyd, 2020 online<br />
<b>Maastrichtian, Late Cretaceous<br />
Lapurr sandstone (= Turkana Grits), Turkana, Kenya<br />
Material</b>- (KNM-WT coll.) (~11-12 m) multiple specimens including cranial material (including
a partial skull - premaxilla, postorbital and braincase), axial material and
appendicular material (including astragalocalcaneum)<br />
<b>Comments</b>-
Discovered in 2004, this is based on unassociated material referred to
one taxon "based on
morphological consistency and on the recovery of specimens from a
narrow stratigraphic and geographic area." Abelisaurid characters
listed by Sertich et al. (2013) include- "a tall, rugose premaxilla, an
anteroventrally inclined posterior border of the postorbital," ... "a
prominent dorsal projection of the parietals and supraoccipital" and an
"ascending process [that] is low and subrectangular, separated from the
anterior surface of the astragalus by a distinct fossa." They
further note the "skull is strongly coossified, with a thickened but
weakly sculptured skull roof", "no prominent cranial ornamentation is
evident" and "the astragalocalcaneum is completely coossified and
displays a prominent transverse sulcus on the anteroventral
surface." The museum collection is based on an azhdarchid
vertebra found by the same team from the same locality (O'Connor et
al., 2011).<br />
Boyd (2020) included the
name Titanovenator kenyaensis in the pdf visual portion of her "Ask a
Geologist" presentation for RUGM. It accompanies a photo of the MCN <span style="font-style: italic;">Carnotaurus </span>cast
that is also labeled Abelisauridae gen. et sp. nov., and based on
the name refers to Sertich et al.'s (2013) giant Kenyan abelisaurid
taxon. It is a nomen nudum as the pdf was not "issued for the
purpose
of providing a public and permanent scientific record" (ICZN Article
8.1.1) and doesn't "state the date of publication in the work itself"
(Article 8.5.2), while the name itself was not "registered in the
Official Register of Zoological Nomenclature (ZooBank)" (Article 8.5.3)
or "accompanied by a description or definition that states in words
characters that are purported to differentiate the taxon" (Article
13.1.1). The name (as "Titanovenator kenyanis") actually predates
Boyd's presentation, originating on a Jurassic Park message board from
2013 in a post under the pseudonym Rex Fan 684, but this is not
considered a source for catalogued nomina nuda here.<br />
<b>References</b>- Sertich, Manthi, Sampson, Loewen and Getty, 2006. Rift Valley
dinosaurs: A new Late Cretaceous vertebrate fauna from Kenya. Journal of Vertebrate
Paleontology. 26(3), 124A.<br />
O'Connor, Sertich and Manthi, 2011. A pterodactyloid pterosaur from the
Upper Cretaceous Lapurr sandstone, west Turkana, Kenya. Anais da
Academia Brasileira de Ciências. 83(1), 309-315.<br />
McCoy, 2013 online. <a href="http://www.jurassicworld3.net/topic/28609">http://www.jurassicworld3.net/topic/28609</a><br />
Sertich, Seiffert and Manthi, 2013. A giant abelisaurid theropod from the Latest
Cretaceous of northern Turkana, Kenya. Journal of Vertebrate Paleontology. Program
and Abstracts 2013, 211.<br />
Boyd, 2020 online. <a href="https://geologymuseum.rutgers.edu/images/geology_museum/events/Ask_A_Geologist_Events/2020_Events/Presentation_Slides/04-30-Melissa-Boyd-Kenya-Geology-Anthro.pdf">Ask a Geologist: Geology & Paleontology of Kenya</a>. Rutgers Geology Museum. 25 pp.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhDu8fQGHLGhqXsWG1-OtFDq97390WYbqAYPzveOCzEfR3F7nkkAcXwXI51eQMl1Tn0fOROnQswpQJV-KR-WOQFlLaYLOULR03CowNT_4JNdJW-r1zb72d0SSz6eWNbZhRmMCtmnD_tB8XALfBq9PCN8qrhI8jeHE2xDABZn27NbE0rkIif8OTREDoVwqQ/s1014/Duwi%20abelisaurid%20teeth.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="583" data-original-width="1014" height="184" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhDu8fQGHLGhqXsWG1-OtFDq97390WYbqAYPzveOCzEfR3F7nkkAcXwXI51eQMl1Tn0fOROnQswpQJV-KR-WOQFlLaYLOULR03CowNT_4JNdJW-r1zb72d0SSz6eWNbZhRmMCtmnD_tB8XALfBq9PCN8qrhI8jeHE2xDABZn27NbE0rkIif8OTREDoVwqQ/s320/Duwi%20abelisaurid%20teeth.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Duwi abelisaurid teeth (MGUP coll.; lost) in (left) labiolingual and (right) basal view (after Gemmellaro, 1921).<br /></td></tr></tbody></table><br /></p><p><b>unnamed Abelisauridae</b> (Di Stefano, 1919)<br />
<b>Late Campanian, Late Cretaceous<br />
Duwi Formation, Al Sharauna, Egypt</b><br />
<b>Material</b>- (MGUP MEGA002; lost?) tooth (16.63x13.01x6.69 mm) (described by Smith and
Lamanna, 2006)<br />
<b>Late Campanian, Late Cretaceous<br />
Duwi Formation, Al Sharauna, Gebel Duwi and/or Gebel Nakheil, Egypt</b><br />(MGUP MEGA001) partial lateral tooth (?x~22.7x? mm) (Di Patti, pers. comm. 6-2023)<br />
(MGUP MEGA002 B) lateral tooth (~35.2x~25x? mm) (Di Patti, pers. comm. 6-2023)<br />
(MGUP MEGA003) partial lateral tooth (?x~30x? mm) (Di Patti, pers. comm. 6-2023)<br />
(MGUP coll.; lost) more than two lateral teeth<br />
<b> Comments</b>- Gemmellaro (1921) referred several remains to <i>Megalosaurus crenatissimus</i>,
from three localities in eastern Egypt. The phosphate layers of Gebel
Duwi are the Late Campanian Duwi Formation (e.g. Salama et al., 2021).
Similarly, "Phosphate mines at Kosseir-el-Khadim, Gebel Nakheil, near
Kosseir [now Quseer], on the Red Sea" are also the Duwi Formation
(Valentine, 1985). The third locality has been mistaken in every
subsequent reference as "Sciarauna-el-Ghibli, presse Sibaiya (Valle del
Nilo)", but the 'c' is actually a poorly photocopied 'e', as indicated
by Di Stefano's (1919) original publication of the expedition which
states "<span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">The
richest and most extensive deposit is the one found on the hill of
Seiarauna, on which are the two villages called Seiarauna-el-Bahri,
i.e. Seiarauna at the river, and Seiarauna-el-Ghibli, i.e. upstream"
(translated). The latter is now known as Ash Sharawinah al Qibliyah,
El-Sharâwna el-Qiblîya or simply Al Sharauna among other variants</span></span></span><span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">, and is indeed directly across the Nile from Sebaiya (also called As Sibaiyyah </span></span></span><span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb"></span></span></span>and
Al Sebaaia West City). Smith and Lamanna (2005) stated the particular
tooth they described was from "near Idfu in the Nile Valley of Egypt
(Fig. 1, exact locality data are unknown; research in Palermo did not
reveal a more accurate location)", but Idfu (also called Edfu) is 30 km
to the southeast. Gemmellaro states "the majority of the specimens
came from [Seiarauna-el-Ghibli]. But a careful research recently
conducted by myself on the abundant material donated by Cortese has
allowed me to find all the types studied also in the locations of Gebel
Nakheil (Kosseir-el-Khadim and Uadi-el-Anz mines) and of Gebel Duwi"
(translated), suggesting no other localities were involved and perhaps
30 km was considered "near" by whoevever wrote that record.<br />
Di Stefano (1919) first reported theropod material from Al Sharauna,
collected by him in 1912 and also donated to the MGUP by Cortese
between then and his publication- "In contact between the phosphates
and the alternating marls, bones and quite a few dinosaurian teeth (<span style="font-style: italic;">Mosasaurus</span>, <span style="font-style: italic;">Platecarpus</span>, <span style="font-style: italic;">Megalosaurus</span>,
etc.) are found, which will be the subject of a special study." That
study was Gemmellaro (1921), who as noted above also discovered
theropod material in the Gebel Duwi and Gebel Nakheil collections.
Unfortunately, Gemmellaro never specifies which of the three localities
any of his discussed specimens is from, nor does he state how many
teeth were in the MGUP collections. Three teeth are figured, with
those in figures 12 and 13 typically abelisaurid with low crowns
serrated to the base both mesially and distally. Curvature and crown
height ratios are most similar to posterior dentary teeth of
<span style="font-style: italic;">Majungasaurus</span>. Gemmellaro compares the teeth favorably to <span style="font-style: italic;">Majungasaurus</span>
syntypes FSL 92.306a and b and a Maevarano tooth figured by Thevenin
1906 (MNHN coll.). A smaller, third tooth is illustrated as
figure 14,
said to be among "some which, although sharing some common features,
differ in the shape of the crown which is not backward curved but is
almost isosceles with the sides having a profile slightly convex or
straight." While Gemmellaro believed "such differences in shape
and in
dimensions between teeth has to be attributed to the diverse locations
they occupied in the jaws, and also to the different age of the
individuals whom the teeth belonged to", the mesiodistally narrower
crown suggests another taxon such as a noasaurid, and that tooth is
listed as Averostra indet. here. Gemmellaro described two unguals
and figured one. While he compares them favorably to <span style="font-style: italic;">crenatissimus</span>
syntype FSL 92.290, the preserved section of the figured specimen is
less tapered in side view with no obvious vascular groove, and this
holds true as well for other <span style="font-style: italic;">Majungasaurus</span> pedal unguals (e.g. FMNH PR 2434, MSNM V6418, V6419) as well as <span style="font-style: italic;">Masiakasaurus</span>
(e.g. FMNH PR 2135) and indeed most theropods. Photos provided by Di
Patti (pers. comm. 6-2023) show the figured element (MGUP MEGA004) is a
sauropod pedal ungual while the other (MGUP MEGA005) compares well to
an ankylopollexian manual ungual I. They are referred to Titanosauria
indet. and Hadrosauroidea indet. here based on age.<br />
Smith and Lamanna (2006) described one of Gemmellaro's teeth in detail
(MGUP MEGA002), although it is not one previously figured. They
identified it as abelisaurid based on "the distal curvature profile
exhibits almost no curvature and is tilted slightly toward the apex in
lateral view" (but this is not true of Gemmellaro's Figure 12), and as
closer to <span style="font-style: italic;">Majungasaurus</span> and Lameta AMNH 1753/1955 than <span style="font-style: italic;">Rugops</span>, <span style="font-style: italic;">Rugops</span>?
sp. UCPC 10 or Bajo Barreal UNPSJB-PV 247 based on "mesial curvature
profile begins at a strong curve at about 1/3 the crown height basal to
the apex (but this is not true of Gemmellaro's Figure 12) and "the ...
interdenticular sulci complex forms an intermediate condition between
being absent and well developed" (unknown in Gemmellaro's figured
specimens). They find "The discriminant analysis (DA) correctly
identified 96.6% of the teeth in the sample and classified MGUP MEGA002
as <span style="font-style: italic;">Majungasaurus</span>" but note "the assignment of the Egyptian tooth to the genus <span style="font-style: italic;">Majungasaurus</span>
in particular is unlikely given that Africa and Madagascar were
separated by the Mozambique Channel throughout the Cretaceous", and
indeed the relatively low sample of abelisaurid teeth tested makes this
questionable. In fact he combination of stout shape plus concave
basal distal edge with a convex apical distal edge is unlike figured <span style="font-style: italic;">Majungasaurus</span> teeth.<br />
Interestingly, the MGUP only has three teeth in their collection now
(Di Patti, pers. comm. 6-2023), none of which are MGUP MEGA002 or match
the three illustrated by Gemmellaro. One (MGUP MEGA002 B) is
abelisaurid based on the slightly convex distoapical edge, although the
mesial edge is similar to Gemmellaro's Figure 12 instead of curving
abruptly at a third of its height. Marginal undulations are
present as in some other abelisaurids (<span style="font-style: italic;">Abelisaurus</span>, <span style="font-style: italic;">Chenanisaurus</span>, <span style="font-style: italic;">Majungasaurus</span>, <span style="font-style: italic;">Skorpiovenator</span>).
There are ~10 mesial serrations per 5 mm and ~11 distal
serrations. The other two teeth (MGUP MEGA001 and MGUP MEGA003)
are more partially preserved but of similar size and serration density
(~10 distal serrations per 5 mm in MEGA001; ~10.5 mesial serrations per
5 mm in MEGA003), and each shows marginal undulations as well, so
they are tentatively placed as Abelisauridae here. There are no
precise locality data for any of them past Maastrichtian of Egypt.<br />
<b>References</b>-
Di Stefano, 1919. Osservazioni sul Cretaceo e sull'Eocene del Deserto
Arabico di el-Sibaiya, nella valle del Nilo. Bollettino del R. Comitato
Geologico d'Italia. Serie V(47), 1-39.<br />
Gemmellaro, 1921. Rettili maëstrichtiani di Egitto. Giornale
di Scienze Naturali ed Economiche. 32, 339-351.<br />
Valentine, 1985. Structure and tectonics of the southern Gebel Duwi
area, Eastern Desert of Egypt. Department of Geology and Geography,
University of Massachusetts. Contribution No. 53, 141 pp.<br />
Smith and Lamanna, 2006. An abelisaurid from the Late Cretaceous of Egypt: Implications
for theropod biogeography. Naturwissenschaften. 93(5), 242-245.<br />
Salama, Altoom, a Allam, Ajarem and Abd-Elhameed, 2021. Late Cretaceous anacoracid sharks (<span style="font-style: italic;">Squalicorax</span>)
from Duwi Formation, Gebel Duwi, central Eastern Desert, Egypt:
Qualitative and quantitative analyses. Historical Biology. 33(11),
3056-3064.<br /><br /><br /><br /><b>Reference</b>- de Souza-Júnior, Candeiro, da Silva Vidal, Brusatte and Mortimer, 2023. Abelisauroidea (Theropoda, Dinosauria) from Africa: A review of the fossil record. Papéis Avulsos de Zoologia. 63, e202363019.</p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com7tag:blogger.com,1999:blog-3248412803814730250.post-15499633138709214942023-05-21T04:28:00.004-07:002023-05-22T00:57:22.233-07:00Raven et al. 2023 on ankylosaur phylogeny missed the shortest trees and should not have dropped Nodosauridae<p>A new paper came out analyzing thyreophoran phylogenetics - Raven et al. (2023). The abstract states "This dataset was analysed using equal- and implied-weights parsimony and Bayesian inference, and further explored using constraint trees and partitioned datasets. Stratigraphical congruence was used to identify a 'preferred tree' and these analyses reveal a novel hypothesis for thyreophoran relationships. The traditional ankylosaurian dichotomy is not supported: instead, four distinct ankylosaur clades are identified, with the long-standing 'traditional' clade Nodosauridae rendered paraphyletic. Ankylosauridae, Panoplosauridae, Polacanthidae and Struthiosauridae have distinct morphotypes..."<br /><br /><b>Four distinct ankylosaur clades with distinct morphotypes?</b><br /><br />It sounds intriguing, but then Polacanthidae as a separate clade or grade has been a viable hypothesis since 1998, so I guess it's the paraphyly of <i>Panoplosaurus </i>and <i>Struthiosaurus </i>versus <i>Ankylosaurus </i>that is new? Looking back, that was ambiguous in Vickaryous et al. (2001), Osi (2005) and Parsons and Parsons (2009) but rejected by Osi and Makadi (2009), Thompson et al. (2012) and Arbour et al. (2016), thus it would be surprising to someone like myself who doesn't specialize in ornithischians so is not all that familiar with character support and such.</p><p></p><p>So let's see what Raven et al. found. They did five basic analyses (A-D with [<b>edited</b> thanks to David Marjanovic's comment] <strike>progressively more</strike> different assumed weighting- unweighted, then k = 3, 8 and 12; E as a Bayesian attempt) then some constraint analyses and ones using only certain parts of the skeleton. I'm going to ignore the latter as I don't think anyone considers "cranial only", "postcranial only" or "armor only" to give better results in Mesozoic dinosaur phylogenetics. In any case, the first analysis (Analysis A) was equal weights, which is the standard for Mesozoic dinosaur analyses and how basically every prior ankylosaur analysis was run. As an aside, statements like "The strict consensus tree (Supplementary material, Fig. S75) shows a lack of resolution in Stegosauria" .... "and Ankylosauria is found in an unresolved polytomy with most stegosaur taxa" just means you haven't pruned enough taxa a posteriori to see the underlying structure. It's annoying the authors never look into what that structure is and leave the tree looking artificially uncertain. But the main result in Ankylosauria is... "There are two clades within Ankylosauria (Ankylosauridae + Nodosauridae)"! So the expected usual result. Hmm. But what about their novel hypothesis of three nodosaur clades at least? <br /><br /><br /></p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhGKG7teX0440n9qBM8t7CqoVCE3QTAphV92QAfhEULXxI5WHE7AbcCaEqd8F5-TEXlRHdgO1-FpXsDh_1toFbwdwA0mh3Vnu2rsQbwTiZPhp0p0FdhPiyelhudmtug515jZXkXMkmC4zYhD-YUprb_yTEm7s7_eYk0BqsY2ucnqDjxltFNU2fr519o/s1260/Raven%20et%20al%202023%20Analysis%201.png" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1208" data-original-width="1260" height="307" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhGKG7teX0440n9qBM8t7CqoVCE3QTAphV92QAfhEULXxI5WHE7AbcCaEqd8F5-TEXlRHdgO1-FpXsDh_1toFbwdwA0mh3Vnu2rsQbwTiZPhp0p0FdhPiyelhudmtug515jZXkXMkmC4zYhD-YUprb_yTEm7s7_eYk0BqsY2ucnqDjxltFNU2fr519o/s320/Raven%20et%20al%202023%20Analysis%201.png" width="320" /></a></div><br /><p></p><p>They state "Within Nodosauridae, there are three groupings of taxa: 'polacanthid' ankylosaurs, but excluding <i>Polacanthus</i>; a 'panoplosaurid' group typified by <i>Edmontonia </i>and <i>Panoplosaurus</i>; and a 'struthiosaurid' group typified by <i>Struthiosaurus </i>and <i>Hungarosaurus</i>." But are there? Just look at the cladogram above (which is only in the supp info). 'Polacanthidae excluding <i>Polacanthus</i>' is <i>Texasetes</i>, the then-unnamed <i>Patagopelta</i>, <i>Sauropelta</i>, <i>Hylaeosaurus</i>, <i>Hoplitosaurus</i>, <i>Tatankacephalus</i>, an undescribed Wessex specimen, <i>Zhejiangosaurus</i>, <i>Antarctopelta </i>and <i>Dongyangopelta</i>. Besides <i>Hoplitosaurus </i>and <i>Hylaeosaurus </i>(which I don't know has ever been recovered as a polacanthid, just guessed to be there), I don't think any of these have been associated with Polacanthidae/inae before. The 'core' polacanthids besides <i>Polacanthus </i>itself are <i>Gastonia</i>, <i>Mymoorapelta </i>and maybe <i>Gargoyleosaurus</i>, so I wouldn't say this clade is reflective of Polacanthidae in any sense. But okay you say, maybe it's not Polacanthidae, but surely the analyses (PLURAL as in the abstract) revealed a 'distinct morphotype' for this <i>Sauropelta</i>-<i>Hylaeosaurus </i>clade of nodosaurs? Looking at their "preferred tree" (Analysis B, k=3) only <i>Patagopelta</i>, <i>Texasetes</i>, <i>Hoplitosaurus </i>and <i>Hylaeosaurus </i>are shared between 'polacanthid' clades, with five new taxa in there compared to Analysis A. And in Analysis C (k=8) only <i>Hoplitosaurus </i>and <i>Texasetes </i>are shared (with seven new taxa), while by Analysis D (k=12) all these taxa are scattered to the wind and there's no equivalent at all. And Analysis E (Bayesian) is one huge polytomy for ankylosaurs. So I would suggest their published results do not support any clade like this that is robust when analyzed under different criteria.<br /></p><p></p><p>But what about Panoplosauridae and Struthiosauridae? First of all, just as there are nine 'nodosaurs' that fall outside the Nodosauridae+ Ankylosauridae split, there are one to five nodosaurids that don't fall into either the panoplosaur or struthiosaur clades. We can't tell how many because Raven et al. do not prune a posteriori to try to resolve any polytomies. But the panoplosaur group is <i>Dracopelta</i>, <i>Aletopelta</i>, both <i>Edmontonia </i>species, <i>Denversaurus </i>and of course <i>Panoplosaurus</i>. The latter three have always been grouped together, but the first two would be interesting if they actually were panoplosaurs. The 'preferred tree' (k=3) takes away <i>Aletopelta </i>and adds <i>Nodosaurus</i>, <i>Anoplosaurus </i>and <i>Tianchisaurus</i>; Analysis C (k=8) has the core three genera plus <i>Dracopelta </i>and <i>Anoplosaurus</i>, while Analysis D (k=12) has everything in A plus <i>Anoplosaurus </i>and <i>Nodosaurus</i>. So <i>Dracopelta </i>is always a panoplosaur and <i>Anoplosaurus </i>is with any unequal weighting, which are the first surprising, new and widely supported nodosaur placements in this study. Yet neither is touted in the text or written up with character support, and honestly the idea of Albian-Cenomanian English <i>Anoplosaurus </i>and especially Jurassic European <i>Dracopelta </i>breaking up not just Campanian-Maastrichtian North American <i>Panoplosaurus</i>+<i>Edmontonia</i>, but the genus <i>Edmontonia </i>itself(!) just seems unlikely. And indeed, a fairly complete <i>Dracopelta </i>specimen (Russo and Mateus, 2023) was recently discovered, and the authors found "<i>D. zbyszewskii</i> [was] consistently recovered as sister taxa of <i>G[argoyleosaurus] parkpinorum</i>, from the Upper Jurassic of Morrison Formation, USA, in a basal ankylosaur group that also includes the other Morrison Formation ankylosaur, <i>M[ymoorapelta] maysi</i>," which matches temporally so much better.<br /><br />As for Struthiosauridae, the unweighted tree would have this include <i>Borealopelta</i>, <i>Minmi</i>, <i>Niobrarasaurus</i>, <i>Polacanthus</i>, <i>Europelta</i>, <i>Liaoningosaurus</i>, <i>Stegopelta</i>, the Paw Paw Formation juvenile (* see below), <i>Hungarosaurus </i>and all three <i>Struthiosaurus </i>species (not monophyletic, at least <i>Tianchisaurus </i>is closer to <i>S. transylvanicus</i> than to <i>S. austriacus</i> or <i>S. languedocensis</i>). The 'preferred tree' (k=3) keeps only <i>Struthiosaurus</i>, <i>Hungarosaurus</i>, the Paw Paw juvenile and <i>Europelta</i>, and adds <i>Silvisaurus </i>and <i>Taohelong</i>. Analysis C (k=8) drops the latter two but adds <i>Tianchisaurus </i>back, and Analysis D (k=12) keeps the two and everything from the unweighted tree plus adds <i>Invictarx </i>and <i>Hoplitosaurus</i>. So far from dividing the 'distinct morphotypes' of Polacanthidae and Struthiosauridae, two of five trees have <i>Polacanthus </i>as a struthiosaur. Here the results are besides the Santonian-Maastrichtian Central European <i>Struthiosaurus</i>/<i>Hungarosaurus</i>, the struthiosaur clade includes <i>Europelta </i>(as guessed by its describers), the Early Cretaceous American Paw Paw juvenile and usually the Jurassic Chinese <i>Tianchisaurus</i>. Yet the authors never mention the Paw Paw juvenile ever falling out here, and have <i>Tianchisaurus </i>as a panoplosaurid because Analysis B is the one time it wasn't the sister to <i>Struthiosaurus transylvanicus</i>. In fact, the authors falsely* state the Paw Paw juvenile is usually a basal ankylosaur and Figure 2 incorrectly* shows it being one in a supposed "Agreement subtree of the three implied weighing analyses (analyses B–D)." How did this happen?! ... <br /><br />(*) I figured it out- the trees in the supp info switched <i>Pawpawsaurus </i>and the Paw Paw juvenile, while figures 1-3 in the paper are correct. So actually their trees have <i>Pawpawsaurus </i>as the struthiosaur sister to <i>Hungarosaurus</i>, which has the same chronostratigraphic issues as the juvenile from the same formation.<br /><br />If we go back to the big picture in Ankylosauria, the unweighted Analysis A gave us a large Nodosauridae with some mostly Jurassic taxa basal to the Nodo-Ankylo split, and a few 'nodosaurs' (<i>Kunbarrasaurus</i>, <i>Peloroplites</i>, <i>Liaoningosaurus</i>) as basal ankylosaurids. 'Preferred' Analysis B (k=3) has polacanthids basal to the split, then panoplosaurs and struthiosaurs sister to each other, so again is pretty standard. Analysis C (k=8) has struthiosaurs, panoplosaurs and polacanthids successively closer to ankylosaurids, so at least that has the paraphyletic nodosaurs the article touts. Finally, Analysis D (k=12) has struthiosaurs further from ankylosaurids than panoplosaurs while polacanthids cease to really exist (their internal specifier <i>Gastonia </i>[see below] is an ankylosaurid but <i>Polacanthus </i>and <i>Hoplitosaurus </i>are struthiosaurs).<br /><br />So I guess if I had a takeaway from their <u>published</u> results (**, see below), it would be that <i>Dracopelta</i> (probably incorrectly) and usually <i>Anoplosaurus </i>are panoplosaurs; <i>Europelta</i>, <i>Pawpawsaurus </i>and usually <i>Tianchisaurus </i>(not even supported by the authors) are struthiosaurs; polacanthids are not strongly supported in any form; and nodosaurs become increasingly paraphyletic with <strike>more weighting</strike> greater values of k (corrected again thanks to David Marjanovic's comment), although you need to get to k=8 for anything really novel. And it's the weighting that is one of my major issues with this paper, because why is k=3 the preferred tree? Because "The stratigraphically most congruent topology, as identified by the four stratigraphical congruence metrics (SCI, RCI, MSM and GER), was Analysis B, and so this was selected as the 'preferred tree'." But if you look at their Table 2 (below), the Bayesian analysis destroys the others at SCI (0.929 vs. 0.438-0.500), but we never get to know what those results are since the authors just leave it as a huge polytomy without further analysis. And in the other three, Analysis B is 16.954, .023 and .005 better respectively, which seems increasingly less important. I have no idea how any of these measures work, but it seems incredibly arbitrary to say whichever k value is best in a majority of four methods wins, ignoring anything quantitative.<br /><br /><br /></p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjS1AjKi9u2KHC1GPmZtDIke91QGUiFe6vMdb4nebfUbRc-EuZafibLYnCA1VO2DCCkwWEyNPYsF3y41CE5XUnO7Z_wgCm3ID1T53llO3Ag7vkKqaV1p2VC9rGyNJpAH3UDTkcuuiptUdqtRUgRiRmoer5v2sNXXkfgldqY6BwieJd0L-Alq2lC8OFM/s477/Raven%20et%20al%202023%20stratcon.png" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="279" data-original-width="477" height="187" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjS1AjKi9u2KHC1GPmZtDIke91QGUiFe6vMdb4nebfUbRc-EuZafibLYnCA1VO2DCCkwWEyNPYsF3y41CE5XUnO7Z_wgCm3ID1T53llO3Ag7vkKqaV1p2VC9rGyNJpAH3UDTkcuuiptUdqtRUgRiRmoer5v2sNXXkfgldqY6BwieJd0L-Alq2lC8OFM/s320/Raven%20et%20al%202023%20stratcon.png" width="320" /></a></div><br /><p></p><p><b>(**) Raven et al. didn't find the shortest trees</b><br /><br />But now we get to the part where I reveal nothing I said above matters, because Raven et al. didn't get the shortest trees. <b>EDIT BELOW</b> <strike>Not even close. Instead of producing "eight MPTs with lengths of 1508 steps", their unweighted matrix Analysis A results in >99999 MPTs of 1464 steps. Here's the real strict consensus with 13 taxa pruned a posteriori for resolution-<br /><br /><br /></strike></p><div class="separator" style="clear: both; text-align: center;"><strike><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg7BN4bJdHVFY9nGXx3j81mdGok_tiJ6oMYJ1Jx4hPpUvSVBa7SayR22eOm4RpbWCeo0mN2Pu4nc37THnfnbvdyVB2iR28tgI8EhHqeg78HPYIzRAy9EAyqEfNgj9gqpCYAmWd9ATlINhG329DlZ33Aj1b76UCRsjxaQCXhk3mAcawOzouhsxwVbCu7/s1309/Raven%20et%20al%202023%20real%20results.png" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1309" data-original-width="898" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg7BN4bJdHVFY9nGXx3j81mdGok_tiJ6oMYJ1Jx4hPpUvSVBa7SayR22eOm4RpbWCeo0mN2Pu4nc37THnfnbvdyVB2iR28tgI8EhHqeg78HPYIzRAy9EAyqEfNgj9gqpCYAmWd9ATlINhG329DlZ33Aj1b76UCRsjxaQCXhk3mAcawOzouhsxwVbCu7/s320/Raven%20et%20al%202023%20real%20results.png" width="220" /></a></strike></div><strike><br /></strike><p><strike>As you can see, it's not the same as Raven et al.'s Figure S75. For one, stegosaurs resolve, although weirdly with <i>Toujiangosaurus</i>+<i>Paranthodon </i>as ankylosaurs. For two it's WAY less resolved in Ankylosauria. All those taxa from <i>Zhejiangosaurus </i>through <i>Ahshishlepelta </i>never form a consistent clade with each other or the four ankylosaur clades, and if you prune all nine genera, struthiosaurs, ankylosaurids and 'panoplosaurs'+polacanthids are still a trichotomy. So their matrix doesn't actually show how these taxa relate (besides polacanthids being sister to 'panoplosaurs'). Isn't it ironic though that we do get Raven et al.'s three nodosaur clades including a classic Polacanthinae that includes <i>Gastonia</i>, <i>Gargoyleosaurus</i> and <i>Mymoorapelta </i>in addition to Jurassic <i>Sarcolestes </i>(and basally some parankylosaurs, but the paper was too late to include <i>Stegouros</i>)? Struthiosauridae is only <i>Struthiosaurus </i>spp. plus <i>Hungarosaurus</i> and <i>Tianchisaurus</i>, which again is funny because the latter was not included in the clade by the authors due only to their Analysis B. 'Panoplosauridae' includes a lot more taxa, and yes those basal ones don't clade with each other or successively to the core group no matter how many are pruned, so that's another real polytomy. As for the pruned taxa-<br /><br /><i>Mongolostegus </i>can at least be sister to <i>Chungkingosaurus </i>or a struthiosaurine.<br /><i>Adratiklit </i>is part of the <i>Dacentrurus</i>+<i>Stegosaurus </i>clade.<br /><i>Anodontosaurus </i>and <i>Scolosaurus </i>are part of the <i>Euoplocephalus</i>+<i>Pinacosaurus </i>clade.<br /><i>Tarchia kielanae</i> is part of the <i>Ankylosaurus</i>+<i>Euoplocephalus </i>clade.<br /><i>Acantholipan </i>can be a 'panoplosaur', a struthiosaurine or outside (by which I mean a taxon closer to <i>Ankylosaurus </i>than <i>Toujiangosaurus </i>but not part of the struthiosaur, ankylosaurid or polacanthid+'panoplosaur' clades shown, though it could be e.g. sister to any of these clades and thus fall under their definitions).<br /><i>Borealopelta </i>can be a struthiosaurine, a 'panoplosaur' or outside(?).<br /><i>Europelta </i>can be a struthiosaurine or outside.<br /><i>Invictarx </i>can be a struthiosaurine, a 'panoplosaur' or outside.<br /><i>Nodosaurus </i>is always closer to <i>Panoplosaurus </i>than <i>Silvisaurus</i>.<br /><i>Patagopelta </i>can be a basal 'panoplosaur' or outside.<br /><i>Pawpawsaurus </i>can be a struthiosaurine or 'panoplosaur'.<br /><i>Stegopelta </i>can be a struthiosaurine or outside.<br /></strike><br /><b>EDIT ADDED 5-21</b>: Thanks to Andrea Cau in the comments for pointing out Raven et al. didn't include their character ordering settings in their txt file. I wrongly assumed it would have a ccode line or a ctype line below the matrix but never scrolled all the way down. I should have been suspicious when I had to manually choose the outgroup instead of them just making <i>Lesothosaurus </i>the first taxon in the matrix. On the one hand, my bad. On the other hand, it's surely best practice to not force your readers to modify the settings of the file you provided to the journal.<br /><br />In any case, it still doesn't matter because Raven et al. STILL didn't find the shortest trees. Instead of producing "eight MPTs with lengths of 1508 steps",
their unweighted matrix Analysis A results in >99999 MPTs of 1506
steps. Here's the real REAL strict consensus with 13 taxa pruned a
posteriori for resolution-<br /><br /><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiKaDlsVUYW8MKX7iLe3sKJBD_leOfZXCOu3H7Hz2rzgL_zfLHiz9lyg_nAAtk0cxOGKbiq6u5b4dQ_PpFRJNnOWuq_0lxluEGt88LEkm0oHKXwO7IOm0Jhtf0C2bo4yxxmrZ82j6MQ_a4F7LfpllLrZnHlVgLKargKK0oiqOclNlcna0bBD2jwu2aN/s1308/Raven%20et%20al%202023%20ordered%20real%20results.png" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1308" data-original-width="1264" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiKaDlsVUYW8MKX7iLe3sKJBD_leOfZXCOu3H7Hz2rzgL_zfLHiz9lyg_nAAtk0cxOGKbiq6u5b4dQ_PpFRJNnOWuq_0lxluEGt88LEkm0oHKXwO7IOm0Jhtf0C2bo4yxxmrZ82j6MQ_a4F7LfpllLrZnHlVgLKargKK0oiqOclNlcna0bBD2jwu2aN/s320/Raven%20et%20al%202023%20ordered%20real%20results.png" width="309" /></a><br /><br />Two steps doesn't sound like much, but it's enough to make 'polacanthids' a grade of basal ankylosaurids, make struthiosaurines nodosaurids as in traditional phylogenies and kick <i>Dracopelta </i>and <i>Anoplosaurus </i>out of panoplosaurs. And yes, <i>Nodosaurus </i>is still closer to <i>Panoplosaurus </i>than <i>Silvisaurus </i>or <i>Struthiosaurus</i>. The toplogy still has <i>Tuojiangosaurus</i>+<i>Paranthodon </i>as ankylosaurs and a struthiosaur <i>Tianchisaurus </i>too.<br /><br /><b>What are these clades named?</b><br /><br />Raven et al. propose new definitions for their three nodosaur families-<br /></p><p>"Panoplosauridae <br />All ankylosaurs more closely related to <i>Panoplosaurus </i>than to <i>Ankylosaurus</i>, <i>Struthiosaurus austriacus</i> or <i>Gastonia burgei<br /></i><br />Polacanthidae <br />All ankylosaurs more closely related to <i>Gastonia burgei</i> than to <i>Ankylosaurus</i>, <i>Panoplosaurus </i>or <i>Struthiosaurus austriacus</i><br /><br />Struthiosauridae <br />All ankylosaurs more closely related to <i>Struthiosaurus austriacus</i> than to <i>Ankylosaurus</i>, <i>Panoplosaurus</i> or <i>Gastonia burgei</i>"<br /><br />Tim Williams has already rightfully complained on the DML that their Polacanthidae definition needs to use <i>Polacanthus foxii</i>. Why is this so hard in 2023?! Do Arbour and the "three anonymous referees" not know the basics of phylogenetic nomenclature? PhyloCode Article 11.10 states "when a clade name is converted from a preexisting name that is typified
under a rank-based code or is a new or converted name derived from the
stem of a typified name, the definition of the clade name must use the
type species of that preexisting typified name or of the genus name from
which it is derived (or the type specimen of that species) as an
internal specifier." We've been complaining about it since Sereno 24 years ago, surely every dinosaur worker knows by now.<br /><br />Another obvious issue is that Nodosauridae has priority over Panoplosauridae, Struthiosauridae and Polacanthidae, so where's Nodosauridae? Raven et al. explain "<i>Nodosaurus </i>is recovered outside of<br />Panoplosauridae in Analyses A and C, further suggesting that application of the name Nodosauridae would add confusion." But as noted above, <i>Nodosaurus </i>is actually always a 'panoplosaur' when Analysis A is run correctly, and <strike>the fact it's supposedly an ankylosaurid sister to <i>Dyoplosaurus </i>when k=8 (fig. S77) should be close to worthless in my view. Unless one of the authors wants to claim whatever character is being weighed eight times more than others could realistically mean this Cenomanian taxon is really within a Campanian species complex that are so similar they were all placed under <i>Euoplocephalus tutus</i> until recently? Also,</strike> <i>Polacanthus </i>doesn't fall in their definition of Polacanthidae in two of their four trees, so why is <u>that</u> still allowed as a family name?<br /><br />In any case, the clades have already been officially defined with PhyloCode registrations, by Madzia et al. (2021). Raven et al. was "Received 7 February 2022", so I don't know why Arbour (a coauthor on Madzia et al.!) or the reviewers would let that stay in the paper. Reading through, Raven et al. actually cite Madzia et al. and state "the underlying philosophy of the latter study is based on the PhyloCode (de Queiroz & Cantino, 2020) and offers an alternative hypothesis to our study, which is framed by the traditional principles of the International Commission on Zoological Nomenclature (1999), and so is not discussed further." Hahahaha I hate to tell you guys, but defining clades based on phylogenetic relationships has nothing to do with the ICZN. And if you were following the "traditional principles" of the 1999 ICZN, you couldn't just throw Nodosauridae away while stating "In the 'preferred' tree Panoplosauridae consists of ... <i>Nodosaurus</i>" and "A clade of generally Late Cretaceous North American taxa is also recovered here and named Panoplosauridae. As well as <i>Denversaurus</i>, <i>Edmontonia </i>spp., <i>Nodosaurus </i>and <i>Panoplosaurus</i>..." Instead you would follow ICZN Article 65.2.3 - "by the discovery that the type genus was, when established,
based on a type species then misidentified, the author may fix as the
type species a nominal species as prescribed in Article 70.3. If the threat cannot be overcome by the fixation of a type species under the provisions of Article 70.3 the case is to be referred to the Commission for a ruling." But that's not happening because I bet they think it's unlikely <i>Nodosaurus </i>is outside Nodosauridae. "... is not discussed further" is short here for "... we know it makes no sense but we don't want to address it."<br /><br />And Madzia et al. do a good job because they actually follow the rules. Except in Raven et al.'s topology Stegosauridae ends up being the <i>Chungkingosaurus</i>+Eurypoda clade due to <i>Huayangosaurus</i>' weird position outside Eurypoda (which supposedly happens in the Bayesian analysis too), which could be saved by adding <i>Ankylosaurus magniventris</i> as an external specifier. <strike>Similarly, Struthiosaurini might benefit from an <i>Ankylosaurus magniventris</i> external specifier due to the polytomy. So here's the actual unweighted results with official clade names-<br /><br /></strike></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><strike><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhdB_TGXND0csACHe1Ijge-OQslQnNwRzkNizANfNHCRRXx0TmP6kM_JaOugvJ5dpVPlDumSFCMpuEAv2eK0f2HabwP3uic_L3gZ7WopmlzeHRP3K_FcxgtCU6xgWAOyQ46nPeqq8R9ewxfbzOCdW8jxAbWxtWiExYBcBxk2mzTrdHb38HNbjEOND6V/s1309/Raven%20et%20al%202023%20real%20results%20with%20clades.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1309" data-original-width="898" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhdB_TGXND0csACHe1Ijge-OQslQnNwRzkNizANfNHCRRXx0TmP6kM_JaOugvJ5dpVPlDumSFCMpuEAv2eK0f2HabwP3uic_L3gZ7WopmlzeHRP3K_FcxgtCU6xgWAOyQ46nPeqq8R9ewxfbzOCdW8jxAbWxtWiExYBcBxk2mzTrdHb38HNbjEOND6V/s320/Raven%20et%20al%202023%20real%20results%20with%20clades.png" width="220" /></a></strike></td></tr><tr><td class="tr-caption" style="text-align: center;"><strike>1. Ankylosauria; 2. Ankylosauridae; 3. Ankylosaurinae; 4. Ankylosaurini;
5. Eurypoda; 6. Huayangosauridae; 7. Nodosauridae; 8. Nodosaurinae; 9.
Polacanthinae; 10. Shamosaurinae; 11. Stegosauria; 12. Struthiosaurini.
Note due to <i>Nodosaurus</i>' possible positions, Panoplosaurini cannot be
placed precisely in this tree.</strike></td></tr></tbody></table><p><br />For a couple final thoughts, Raven et al.'s definition for Shamosaurinae ("All ankylosaurid ankylosaurs more closely related to <i>Shamosaurus </i>than to <i>Ankylosaurus</i>") is better than the official definition ("max ∇ (<i>Gobisaurus domoculus</i> Vickaryous et al., 2001 & <i>Shamosaurus scutatus</i> Tumanova, 1983 ~ <i>Ankylosaurus magniventris</i> Brown, 1908") because <i>Gobisaurus </i>has no reason to be involved. Also, Kirkland et al.'s (2013) definition of Struthiosaurinae is just horrible- "the most inclusive clade containing <i>Europelta </i>but not <i>Cedarpelta</i>, <i>Peloroplites</i>, <i>Sauropelta </i>or <i>Edmontonia</i>." Doesn't separate it from ankylosaurids or polacanthines, includes <i>Cedarpelta </i>and <i>Peloroplites </i>which have no agreed upon positions or competing family-level names, AND doesn't use <i>Struthiosaurus austriacus</i> as the internal specifier. Oof. And why have "Bayesian analysis' as a keyword in Raven et al. (2023), run the Bayesian analysis for a week, mention it in the abstract, and despite having by far the best SCI score just leave it as a huge ankylosaur polytomy that has no effect on its conclusions?<br /><br /><b>So does nobody test run these analyses before they are accepted?</b><br /><br />I began this post just planning to harp on peoples' seeming inability to learn Phylocode Article 11.10, but it ended up so much worse. It's not like I searched and scoured to find a better cladogram in the data, TNT popped it out in (checking) under 30 seconds. And then there's the <i>Pawpawsaurus </i>vs. Paw Paw juvenile switch that made the supp info wrong that I guess nobody noticed? I'm just a person looking over the article on a random Saturday, not a journal editor or a professional whose job description woefully includes peer reviewing. And sure mistakes are made, maybe the authors are anti-Phylocode despite proposing phylogenetic definitions (what organization is going to lend yours any validity?) and maybe nobody on the team knows how to use TNT, but I also just found the basic results quote from the top of my post to be misleading. The published results did NOT show that nodosaurs fall into three "distinct morphotypes" past the standard Latest Cretaceous Central European struthiosaurines and North American panoplosaurins except for maybe 2-5 additional genera, NOR that they were more paraphyletic than normally thought and basically nothing was shown about polacanthines, with the statement "'polacanthid' ankylosaurs, but excluding <i>Polacanthus"</i> in Analysis A being plain wrong. My best guess is that after getting pretty inconclusive results the team went with the "Nodosauridae isn't real" gimmick despite the hypocrisy in keeping Polacanthidae. I think the real message of the analysis (assuming accurate scorings) is that <strike>ankylosaurs need a lot more characters analyzed to determine their basal relationships.</strike> the basic topology from twenty years ago is still most parsimonious.<br /><br /><br /><b>References</b>- Vickaryous, Russell, Currie and Zhao, 2001. A new ankylosaurid (Dinosauria:<br />Ankylosauria) from the Lower Cretaceous of China, with comments on ankylosaurian relationships. Canadian Journal of Earth Sciences. 38(2), 1767-1780.<br /><br />Osi, 2005. <i>Hungarosaurus tormai</i>, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary. Journal of Vertebrate Paleontology. 25(2), 370-383.<br /><br />Osi and Mak adi, 2009. New remains of <i>Hungarosaurus tormai</i> (Ankylosauria, Dinosauria) from the Upper Cretaceous of Hungary: Skeletal reconstruction and body mass estimation. Palaontologische Zeitschrift. 83, 227-245.<br /><br />Parsons and Parsons, 2009. A new ankylosaur (Dinosauria: Ankylosauria) from the Lower Cretaceous<br />Cloverly Formation of central Montana. Canadian Journal of Earth Sciences. 46(10), 721-738.<br /><br />Thompson, Parish, Maidment and Barrett, 2012. Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora). Journal of Systematic Palaeontology. 10, 301-312.<br /><br />Kirkland, Alcal a, Loewen, Esp ılez, Mampel and Wiersma, 2013. The basal nodosaurid ankylosaur <i>Europelta carbonensis</i> n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of northeastern Spain. PLoS ONE. 8, e0080405.<br /><br />Arbour, Zanno and Gates, 2016. Ankylosaurian dinosaur palaeoenvironmental associations were influenced by extirpation, sea-level fluctuation, and geodispersal. Palaeogeography, Palaeoclimatology, Palaeoecology. 449, 289-299.<br /><br />Madzia, Arbour, Boyd, Farke, Cruzado-Caballero and Evans, 2021. The phylogenetic nomenclature of ornithischian dinosaurs. PeerJ. 9, e12362.<br /><br />Raven, Barrett, Joyce and Maidment, 2023. The phylogenetic relationships and evolutionary history of the armoured dinosaurs (Ornithischia: Thyreophora). Journal of Systematic Palaeontology. 21(1), 2205433.<br /><br />Russo and Mateus, 2023. Review of <i>Dracopelta zbyszewskii</i>, an ankylosaur from the Upper Jurassic of Portugal. 14th Symposium on Mesozoic Terrestrial Ecosystems and Biota. The Anatomical Record. 306(supp. 1), 221-223.</p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com26tag:blogger.com,1999:blog-3248412803814730250.post-67189629389800015802023-01-20T07:40:00.003-08:002023-01-20T23:57:11.179-08:00Are uncinate processes ancestral to Archosauria? A look at Wang et al 2023<p align="JUSTIFY" style="margin-bottom: 0in; text-indent: 0.5in;">A paper was published on January 17th (Wang et al., 2023) that I've been looking forward to since its SVP 2020 poster (Wang et al., 2020). The basic idea is that uncinate processes leave attachment scars on ribs even if the processes themselves are unossified, and these scars show that uncinate processes were much more widespread than the record of ossified processes would suggest. In theropods, ossified uncinate processes are only known in many pennaraptorans, <i>Pelecanimimus </i>and the noasaurid "Sidormimus" (Sereno, 2010; not mentioned by Wang et al., 2023) for instance, but Wang et al. found scars in <i>Struthiomimus</i>, <i>Gorgosaurus </i>and <i>Allosaurus</i>, all of which definitely lacked ossified processes. Similarly, Wang et al. identified scars in <i>Apatosaurus</i>, when I don't think any sauropodomorph has been reported with ossified processes.<br /><br />So my first question in 2020 was "do any archosaurs NOT show these scars?", and the SVP abstract and poster didn't say. Even in the published paper, the most we get in the main section is "Using an alternate coding approach in which uncinate processes were coded as absent in taxa represented by five or more dorsal vertebral ribs that all lacked uncinate scars, nine archosaur taxa were coded as lacking uncinate processes." Which nine taxa? Who knows. The Methods section at the end of the paper similarly says "Our alternative coding approach, which was used to test the stability of the results obtained under our preferred coding approach, differed in that uncinate processes were coded as absent in taxa for which at least five vertebral ribs were available, regardless of their state of preservation, and showed no sign of uncinate processes or uncinate scars. This resulted in coding uncinate processes as absent in nine taxa." It does detail that their outgroup, the proterochampsian <i>Chaneresuchus</i>, lacks the scars so was assumed to lack any uncinate processes, but that's it.<br /><br />Their only figure besides two showing the scars is Figure 3, an "informal consensus cladogram", whose caption reads "Major clades of Archosauria with evidence of cartilaginous uncinate processes are labelled and shaded blue; clades with evidence of ossified uncinate processes are labelled and shaded pinkish red; clade with evidence of both cartilaginous and ossified uncinate processes is labelled and shaded purple; and clades for which no evidence is available are labelled and shaded grey." Clades shaded gray are Shartegosuchidae, Pachycephalosauria, Macronaria and Megalosauroidea, the latter being the most basal theropod clade shown. I guess the nine mystery taxa belong to those clades? Also note every branch in each blue clade is blue, every branch in every red clade is red, etc., implying homogeniety. Here's the figure-</p><p align="JUSTIFY" style="margin-bottom: 0in; text-indent: 0.5in;"> </p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqYO-HMfVGJdJKJ2xp4y-c2XbrAlxtyHM6n2pYcFLHUsLjtVaKKSUX67VfEImUps2c3Eh2blLQ3i9bibeTGJ_8FhzAPp2LOTL_3kjR5xmtJM28GPmYO-SaO3HwkjDAl2OxyjeM0cyim2E34GznEDtNj-ztdQRMi5cL18elfcTWCa8l8gS3hpjvx43f/s1070/Wang%20et%20al%20Fig3.png" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1070" data-original-width="1000" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqYO-HMfVGJdJKJ2xp4y-c2XbrAlxtyHM6n2pYcFLHUsLjtVaKKSUX67VfEImUps2c3Eh2blLQ3i9bibeTGJ_8FhzAPp2LOTL_3kjR5xmtJM28GPmYO-SaO3HwkjDAl2OxyjeM0cyim2E34GznEDtNj-ztdQRMi5cL18elfcTWCa8l8gS3hpjvx43f/s320/Wang%20et%20al%20Fig3.png" width="299" /></a></div><p align="JUSTIFY" style="margin-bottom: 0in; text-indent: 0.5in;"><br />It turns out the identity of the Mystery Nine is in Table 2 of the Supplementary Information, under Alternate Coding. In the preceding paragraph they again report "Under
the alternate coding method, nine taxa were scored as lacking
uncinate processes." These taxa are- <i>Lotosaurus</i>, <i>Protosuchus</i>, <i>Lagerpeton</i>, <i>Plateosaurus</i>, <i>Anchisaurus</i>, <i>Camarasaurus</i>, <i>Camptosaurus</i>, <i>Parasaurolophus</i>, <i>Edmontosaurus</i>, <i>Protoceratops </i>and <i>Chasmosaurus</i>. Wait, that's actually eleven taxa, not nine, despite the number nine being stated three times. On the one hand, note none of these are shartegosuchids, pachycephalosaurs, or megalosauroids, so why are those clades even in the cladogram?! On the other hand, note that <i>Lotosaurus </i>and <i>Protosuchus </i>would make two gray nodes basal to mesoeucrocodylians, <i>Lagerpeton </i>would be a gray node basal to dinosaurs, and <i>Plateosaurus </i>and <i>Anchisaurus </i>would be two extra gray nodes basal to sauropods. Also all of the Ornithopoda and Ceratopsida [sic] branches are blue when they shouldn't be. So the figure is putting in artificial discrepancy and leaving out most of the actual discrepancy.<br /><br />The big methodological problem with the paper is that their Preferred Coding approach scored "the
presence of uncinate processes as uncertain (?) if evidence of
uncinate processes or scars was lacking." So the Mystery "Nine" were actually all scored as unknown, even though <i>Chaneresuchus </i>was scored as absent (0) based on identical evidence. So obviously if you score all the archosaurs as present or unknown and the sole non-archosaur as absent, you're going to get archosaurs ancestrally having uncinate processes. And when they use the Alternate Coding of taxa without scars not having uncinate processes, they themselves report- <br /><br />"Maximum likelihood and Bayesian inference recovered cartilaginous uncinate processes as the most likely condition at Archosauria (pml=0.61, pmb=0.55), but only when branch length estimates were incorporated. By contrast, maximum likelihood excluding branch length estimates recovered the absence of uncinate process as the most likely condition at Archosauria (pml=0.92). The ancestral condition at Dinosauria could not be recovered with confidence using either maximum likelihood or Bayesian inference (pml=0.57, pmb ≈ 0.33). Maximum likelihood recovered ossified uncinate processes as the most likely condition at Maniraptoriformes (pml=0.90) and Pennaraptora (pml=0.99), but only when branch length estimates were excluded from the analysis. Bayesian inference could not confidently recover the ancestral conditions at Maniraptoriformes (pmb=0.33) and Pennaraptora (pmb=0.33)."<br /></p>
<p>So not strongly supporting "deep reptilian evolutionary roots of a major avian respiratory adaptation", and that's not even going into how the only non-avian paravians they scored were four dromaeosaurids (<i>Microraptor</i>, <i>Saurornitholestes</i>, <i>Velociraptor </i>and <i>Deinonychus</i>) with ossified processes, when at least <i>Archaeopteryx</i>, anchiornithines and omnivoropteryids lacked them, so anything about Pennaraptora's ancestral state is going to need more than "oviraptorosaurs, dromaeosaurids, <i>Rhea</i>, <i>Gallus </i>and <i>Lithornis</i> - yes, <i>Chauna </i>- no."<br /><br />And yeah, the living bird <i>Chauna </i>has no uncinate processes and lacks any scars for them, so was rightfully scored 0. Yet fossil taxa with the same morphology were scored unknown. Which again points to the flaw in their Preferred Coding, but also suggests we might expect homoplasy in other parts of Archosauria as well. So lambeosaurines and edmontosaurins could have lacked uncinates while kritosaurins had them, following the osteological evidence, for instance.<br /><br />In conclusion, it's an excellent idea to look for osteological correlates to unossified uncinate processes, and we got some real data hidden in the supplementary information, but any use was marred by unforced errors like a misleading main figure and a nonsensical scoring methodology. I wouldn't doubt uncinates were primitive to at least averostrans, given "Sidormimus" and <i>Edmarka </i>(check Figure 14A of Bakker et al., 1992; so Megalosauroidea should have ironically been blue), but the concept deserved better vetting than this.<br /><br /><b>References</b>- Bakker, Kralis, Siegwarth and Filla, 1992. <i>Edmarka rex</i>,
a new, gigantic theropod dinosaur from the Middle Morrison Formation, Late Jurassic
of the Como Bluff outcrop, with comments on the evolution of the chest region
and shoulder in theropods and birds and a discussion of the five cycles of originn
and extinction among giant dinosaurian predators. Hunteria. 2(9), 1-24.<br /><br />Sereno, 2010. Noasaurid (Theropoda: Abelisauroidea) skeleton from Africa shows
derived skeletal proportions and function. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 162A.<br /><br />Wang, Sullivan and LeBlanc, 2020. Anatomical and histological data indicate uncinate processes to be homologous across Archosauria. Journal of Vertebrate Paleontology. Program and Abstracts 2020, 334.<br /><br />Wang, Claessens and Sullivan. 2023. Deep reptilian evolutionary roots of a major avian respiratory adaptation. Communications Biology. 6:3.<br /><br /><br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com2tag:blogger.com,1999:blog-3248412803814730250.post-73438981423738481332023-01-09T02:38:00.000-08:002023-01-09T02:38:36.640-08:00"Scrotum humanum" a torvosaur and Jurassic Chinese theropod updates - The Theropod Database January 2023<p>Happy New Years everyone! Sorry about how long it took to post the <a href="https://www.theropoddatabase.com/Updates.htm" target="_blank">update</a>, but I was covering too many sections at once.<br /><br /><a href="https://tetzoo.com/blog/2022/12/12/robert-plots-lost-dinosaur-bone" target="_blank">Darren Naish's post</a> on Middle Jurassic British theropod femur OUMNH J29757 and "Scrotum humanum" led me to do something that's been on my list for a while- compare the one original figure of "Scrotum" to other theropods to see if is really Megalosaurus as has long been assumed. Turns out it's closer to another closely related taxon...<br /><br /><br /><b>"Scrotum"</b> Brookes, 1763<br />
<b>"S. humanum"</b> Brookes, 1763<br />
<b>Aalenian-Bajocian?, Middle Jurassic<br />unknown quarry, Inferior Oolite?, Cornwell, England </b><br />
<b>Material</b>- (lost) distal femur (~235 mm trans)<br />
<span style="font-weight: bold;">Diagnosis</span>- (proposed) ectocondylar tuber limited to medial half of ectocondyle; ectocondyle subequal in size and shape to endocondyle.<br />
<b>Comments</b>-
Originally described and illustrated by Plot (1677) as the distal femur
of a giant human, this is famous as being the first Mesozoic dinosaur
bone to be published. Brookes (1793) later summarized Plot's
description and opinions but labeled the specimen <span style="font-style: italic;">Sctrotum Humanum</span>
in his plate. Rieppel (2022) notes that "At the top of page 317,
Brookes (1763) noted that 'other stones have been found exactly
representing the private parts of a man; and others in the shape of
kidneys . . . ', and continued further down on the same page" described
the femur. As "the plates and the individual figures they contain are
not numbered separately, but are identified by the pagination number of
the page on which the respective specimens are mentioned or described.
The conclusion seems to be that the illustrator took the femur fragment
to be an example of those stones referred to on page 317 as 'exactly
representing the private parts of a man', and erroneously labelled it
accordingly." Phillips (1871) believed it was from the Inferior Oolite
(Aalenian-Bajocian) and stated "It may have been the femur of a large
megalosaurus or a small ceteosaurus" without evidence. As described by
Delair and Sargeant (1975), Halstead (1970) "pointed out that because
of its date of publication (post-Linnean, i.e. after 1758), this
binomen can be considered a perfectly valid publication of the first
generic and specific name ever applied to dinosaurian remains. It is
perhaps fortunate that the name was not thereafter employed by any
subsequent worker, and thus Scrotum humanum Brookes must be treated as
a nomen oblitum and discarded." They (and Halstead) considered it more
probable to be <span style="font-style: italic;">Megalosaurus</span> than <span style="font-style: italic;">Cetiosaurus</span>
without evidence and stated "The specimen unfortunately is lost."
Halstead and Sarjeant (1993; publication duplicated in 1995) noted that
while Scrotum should be treated as a nomen oblitum under ICZN Article
23b (First Edition), "no application was made then, or has been made
since, for the formal suppression of Brookes's binomen." The Third
Edition of the ICZN came out in 1985 and eliminated the nomen oblitum
clause, so the authors petitioned the ICZN in 1992 "(I) to use its
plenary powers to suppress the generic name <span style="font-style: italic;">Scrotum</span> Brookes, 1763 and the specific name <span style="font-style: italic;">S. humanum</span> Brookes, 1763; (2) to retain on the Official List of Generic Names in Zoology the name <span style="font-style: italic;">Megalosaurus</span> Buckland in Parkinson, 1822, type species by subsequent designation <span style="font-style: italic;">M. bucklandi</span> Meyer. 1832. (3) to retain on the Official List of Specific Names in Biology the name <span style="font-style: italic;">bucklandi</span> as published in the binomen <span style="font-style: italic;">Megalosaurus bucklandi</span> (specific name of the type species of <span style="font-style: italic;">Megalosaurus</span>
Buckland in Parkinson, 1822, by designation in Meyer, 1832); (4) To
place on the Official List of Rejected and Invalid Generic Names in
Zoology the name <span style="font-style: italic;">Scrotum</span> Brookes, 1763; (5) to place on the Official List of Rejected and Invalid Specific Names in Zoology the name <span style="font-style: italic;">humanum</span> Brookes, 1763, as published in the binomen <span style="font-style: italic;">Scrotum humanum</span>,
and as suppressed in (1) above." as listed in 1993. As recalled by the
authors, Tubbs (Executive Secretary to the ICZN) replied later that
year that "The text on p. 301 of Brookes (1763) makes it quite clear
that the two words "Scrotum humanum" on the plate were a description of
a specimen, and that Brookes did not establish a genus <span style="font-style: italic;">Scrotum</span> or a species <span style="font-style: italic;">humanum</span> (any more than he did a species <span style="font-style: italic;">Kidney stone</span> on the same plate!). The words just happened to be Latin." Furthermore, since "[the name S<span style="font-style: italic;">crotum humanum</span>]
has never been used as a scientific name", it "is therefore unavailable
under Article 11d of the Code" (Third Edition- "Names to be treated as
valid when proposed. - Except as in (i) below, a name must be treated
as valid for a taxon when proposed unless it was first published as a
junior synonym and subsequently made available under the provisions of
Section e of this article."). Finally, because "Plot's long-lost
specimen was ... not certainly, a <span style="font-style: italic;">Megalosaurus</span>
bone", Tubbs wrote that "the Commission is willing to take action only
when there is an appreciable and real, as opposed to hypothetical,
threat to stability or nomenclature. This is not the case for <span style="font-style: italic;">Megalosaurus</span>." Note that Tubbs was incorrect that Brookes ever specified <span style="font-style: italic;">Scrotum Humanum</span>
was a description instead of a name, with page 301 being an unrelated
section on plant fossils, so his use of Article 11d was unwarranted
although recently supported by Rieppel's logic. He was also wrong
that it had never been used as a scientific name, as Molnar et al.
(1990) listed <span style="font-style: italic;">Scrotum humanum</span> as a carnosaur nomen dubium. Under the current
ICZN, "Scrotum humanum" would be a nomen nudum based on Article 11.5-
"To be available, a name must be used as valid for a taxon when
proposed." Tubbs was right that the referral to <span style="font-style: italic;">Megalosaurus</span> was merely hypothetical though, as it has never been supported by published evidence and seems unwarranted.<br />
The femur is dissimilar from <span style="font-style: italic;">Cetiosaurus</span>
(both the lectotype OUMNH J13615 and the Rutland specimen LCM
G468.1968) in being 45-86% larger, having a distally extended medial
condyle, and a fibular groove placed at the lateral edge. Note the
estimated transverse diameter is based on Plot's statement the
narrowest shaft circumference was 15 inches (= 381 mm) and scaled from
the figure. Compared to this, <span style="font-style: italic;">Megalosaurus</span>
femora are slightly smaller (shaft diameter 265-343 mm), and differ in
having a more distomedially extended and pointed medial condyle, more
laterally positioned ectocondylar tuber, and a straight lateral edge
until the distal extent of the tuber. These same differences are also
usually present in e.g. <span style="font-style: italic;">Cruxicheiros</span>, piatnitzkysaurids, <span style="font-style: italic;">Eustreptospondylus</span>, <span style="font-style: italic;">Erectopus</span>, <span style="font-style: italic;">Allosaurus</span>, and <span style="font-style: italic;">Juratyrant</span>
among large Jurassic theropods whose distal femora are undistorted and
figured in posterior view. If "Scrotum" is from the Inferior Oolite it
is also earlier than <span style="font-style: italic;">Megalosaurus</span>, and differs from the contemporaneous <span style="font-style: italic;">Magnosaurus</span>
in the same ways when preserved (more laterally positioned ectocondylar
tuber; straight lateral edge until the distal extent of the tuber),
although it could derive from <span style="font-style: italic;">Duriavenator</span> with which it cannot be compared. <span style="font-style: italic;">Sinraptor dongi</span>
is slightly more similar to "Scrotum" in having a convex lateral edge
alongside the ectocondylar tuber, but it is "Brontoraptor" which is
most similar in having that character, an evenly rounded medial condyle
and a more medially placed ectocondylar tuber. <span style="font-style: italic;">Torvosaurus</span>
(ML 632) shows the last character at least but cannot be evaluated for
the rest, so "Scrotum" might be best characterized as a torvosaur and
may relate to <span style="font-style: italic;">"Megalosaurus"</span> "phillipsi" from the Kimmeridgian of England that also has characters similar to "Brontoraptor" and <span style="font-style: italic;">Torvosaurus</span>.
"Brontoraptor" also has a similar circumference (376 mm) and internal
cavity size based on Siegwarth et al.'s Figure 8E. Whether the
remaining differences (ectocondylar tuber limited to medial half of
ectocondyle; ectocondyle subequal in size and shape to endocondyle) are
genuine or illustration error caused by Plot generalizing then
unfamiliar megalosauroid anatomy is uncertain. Note "Scrotum" can be
excluded from Ceratosauria based on the absence of a tall anteromedial
crest, and from Coeluridae, Proceratosauridae and Maniraptoromorpha
based on the deep extensor groove described by Plot and large size
(with occasional exceptions, e.g. <span style="font-style: italic;">Yutyrannus</span>).<br />
<b>References</b>- Plot, 1677. The Natural History of Oxford-shire, being an
essay towards the Natural History of England. Oxford. 358 pp.<br />
Brookes, 1763. The Natural History of Waters, Earths, Stones, Fossils and Minerals,
with their Virtues, Properties and Medicinal Uses: To which is added, the methods
in which Linnaeus has treated these subjects. Vol. 5. J. Newberry. 364
pp.<br />
Robinet, 1768. Vue philosophique de la gradation naturelle des formes
de l'être, ou les essais de la nature qui apprend a faire l'homme.
Harrevelt. 260 pp.<br />
Phillips, 1871. Geology of Oxford and the Valley of the Thames. Oxford at the
Clarendon Press. 523 pp.<br />
Halstead, 1970. <i>Scrotum humanum</i> Brookes 1763 - the first named dinosaur.
Journal of Insignificant Research. 5(7), 14-15.<br />
Delair and Sargeant, 1975. The earliest discoveries of dinosaurs. Isis. 66, 5-25.<br />
Buffetaut, 1979. A propos du reste de dinosaurien le plus anciennement
décrit: l'interprétation de J.-B. Robinet (1768). Histoire et Nature.
14, 79-84.<br />
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 169-209.<br />
Halstead and Sarjeant, 1993. <i>Scrotum humanum</i> Brookes - the earliest name
for a dinosaur? Modern Geology. 18, 221-224.<br />
Halstead and Sarjeant, 1995. <i>Scrotum humanum</i> Brookes - the
earliest name for a dinosaur? In Sarjeant (ed.), 1995. Vertebrate
Fossils and the Evolution of Scientific Concepts; A tribute to L.
Beverly Halstead. Gordon and Breach. 219-222.<br />
Delair and Sargeant, 2002. The earliest discoveries of dinosaurs: The
records re-examined. Proceedings of the Geologists' Association. 113,
185-197.<br />
Rieppel, 2022 (online 2021). The first ever described dinosaur bone
fragment in Robinet's philosophy of nature (1768). Historical Biology.
34(5), 940-946.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiuBmqEPhU6-YMbpOLUx6mz65s6WJSSAGvOS6dd85mKaItK9Ir9Eb7SNZILC4TFDMmttWD0-EzNdiITna8SJZfuOQKFIrw2MULEBJIVg2FsKeGGxlD6HG_alyO-PxtZFNpwTeLnLyOC2SEom9p-A_1wkUXmka-EtF3ELYYsIOj3NAjOQd-hn0eAS9Rl/s1866/Scrotum%20vs%20other%20megalosaurids.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="587" data-original-width="1866" height="101" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiuBmqEPhU6-YMbpOLUx6mz65s6WJSSAGvOS6dd85mKaItK9Ir9Eb7SNZILC4TFDMmttWD0-EzNdiITna8SJZfuOQKFIrw2MULEBJIVg2FsKeGGxlD6HG_alyO-PxtZFNpwTeLnLyOC2SEom9p-A_1wkUXmka-EtF3ELYYsIOj3NAjOQd-hn0eAS9Rl/s320/Scrotum%20vs%20other%20megalosaurids.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">"Scrotum humanum" distal femur (lost; leftmost; after Plot, 1677), compared to (left to right) "Brontoraptor" (TATE 0012; after Siegwarth et al., unpublished), <i>Megalosaurus bucklandii</i> (NHMUK 31806; after Benson, 2010b), <i>Magnosaurus nethercombensis</i> (OUMNH J12143; after Benson, 2010a).<br /></td></tr></tbody></table><br /><p>Also updated is <i>Chienkosaurus</i>, which thanks to the recent description of <i>Sinraptor dongi</i> teeth by Hendrickx et al. (2020), I feel can be placed in Metriacanthosauridae. This and <i>Szechuanosaurus </i>were coincidentally recently reviewed by Curtice at his blog I've started following, <a href="https://www.fossilcrates.com/blogs/news" target="_blank">Dr. BC's Hindsight at Fossil Crates</a>, which I highly recommend.<br /><br /><b><i><a name="Chienkosaurusceratosauroides"></a>Chienkosaurus</i></b> Young,
1942<br />
<i><b>C. ceratosauroides</b></i> Young, 1942<br />
<b>Tithonian?, Late Jurassic<br />IVPP locality 47, upper Guangyuan Group, Sichuan, China</b><br />
<b>Lectotype</b>- (IVPP V237A) (~8 m) posterior premaxillary tooth (44x16x12 mm)<br />
<b>Referred</b>- ?(IVPP V193) ulna (164 mm) (Young, 1942)<br />
<b>Bathonian-Callovian?, </b><b>Middle Jurassic<br />IVPP locality 49, middle Guangyuan Group, Sichuan, China</b><br />
?(IVPP V190) (~5 m) ~ninth caudal centrum (66 mm) (Young, 1942)<br />
<span style="font-weight: bold;">Other diagnoses</span>- Young (1942) originally diagnosed <span style="font-style: italic;">Chienkosaurus</span>
with- "Teeth thick and sharply pointed with fine palisade
denticulations on both sides. The anterior which are finer than the
posterior ones push lingually<br />
towards the base and form a ridge topping at a distance before the base of the tooth."<br />
<b>Comments</b>-
The material was discovered in late Spring 1941, with the type
consisting of four isolated teeth IVPP V237A-D. Young's (1942)
diagnosis was "Mainly based upon" the largest tooth (V237A), with the
three smaller teeth considered immature and (possibly incorrectly)
lacking their bases. He stated "The general shape of the teeth
resembles that of <span style="font-style: italic;">Labrosaurus stechowi</span>" which was prescient as both are based on mesial dentition, and considered <span style="font-style: italic;">Chienkosaurus</span> a ceratosaurid based on the questionably referred postcrania. Ironically, <span style="font-style: italic;">"Labrosaurus" stechowi</span> is now thought to be ceratosaurid, but as Young noted <span style="font-style: italic;">Chienkosaurus</span> lacks its lingual fluting which has proven to be a ceratosaurid character. Subsequently, <span style="font-style: italic;">Chienkosaurus</span>
was generally placed in Megalosauridae (e.g. Romer, 1956; Steel, 1970;
Dong et al., 1978) when it was used as a waste basket for almost all
large Jurassic theropods including <span style="font-style: italic;">Ceratosaurus</span> and later <span style="font-style: italic;">Yangchuanosaurus</span>. Note Huene (1959) when citing <span style="font-style: italic;">Chienkosaurus</span> as named in 1958 from the Late Cretaceous of Shantung meant to list <span style="font-style: italic;">Chingkankousaurus</span>. Dong et al.
(1983) reported that "Rozhdestvensky (1964) proposed that the four
teeth of <span style="font-style: italic;">Chienkosaurus</span> could
possibly belong to the Crocodilia" (translated), but which work this
corresponds to was not listed in the bibliography and cannot be determined. Dong et al.
also stated "during the editing of "The Handbook of Chinese Fossil
Vertebrates," Zhiming Dong conducted a review of these four specimens
and formally confirmed that the best preserved tooth among the V237
collection was a premaxillary tooth of a carnosaurian dinosaur, but
that the remaining three teeth were assignable to the crocodile <span style="font-style: italic;">Hsisosuchus</span>." The dentition of <span style="font-style: italic;">Hsisosuchus</span>
has not been described or figured in enough detail to distinguish it
from theropods, but two of the teeth (IVPP V237B and V237D)
are similar in being short and barely recurved with a high crown base
ratio, characters shared with the tooth figured separately in
<span style="font-style: italic;">Hsisosuchus</span>' type description. They are provisionally placed in <span style="font-style: italic;">Hsisosuchus</span> sp. here. The third supposed <span style="font-style: italic;">Hsisosuchus</span>
tooth (IVPP V237C) is different in having a distinctly D-shaped section
with strong carinae somewhat like Guimarota tyrannosauroid premaxillary
tooth IPFUB GUI D 89, so is provisionally placed in Tyrannosauroidea
here. Retaining only one of <span style="font-style: italic;">Chienkosaurus</span>'
syntype teeth in the genus would make it the lectotype, and as ICZN
Article 74.5 states "In a lectotype designation made before 2000,
either the term "lectotype", or an exact translation or equivalent
expression (e.g. "the type"), must have been used or the author must
have unambiguously selected a particular syntype to act as the unique
name-bearing type of the taxon", and Dong et al. explicitly make <span style="font-style: italic;">Chienkosaurus</span> a synonym of <span style="font-style: italic;">Szechuanosaurus</span>, and of <span style="font-style: italic;">Chienkosaurus</span>'
syntypes only consider IVPP V237A to be theropodan, this is here
considered a valid lectotype designation. Rozhdestveksy (1977) earlier
listed <span style="font-style: italic;">Szechuanosaurus campi</span> and <span style="font-style: italic;">Chienkosaurus ceratosauroides</span> as "synonyms?" in his Table 1 without comment, while Dong et al.'s synonymization was based on examining <span style="font-style: italic;">Yangchuanosaurus</span>
teeth from CV 00214 to correctly determine "the differences among
carnosaur dentitions are due only to being in a different position in
the dentition" and noting <span style="font-style: italic;">Chienkosaurus</span>' and <span style="font-style: italic;">Szechuanosaurus</span>'
types are from the same locality. While this indeed makes it possible
they even derive from the same individual, none of the teeth have been
shown to be diagnostic within metriacanthosaurids, and synonymization
should be based on autapomorphies or unique combinations of characters
instead of provenance. This synonymization of part of the <span style="font-style: italic;">Chienkosaurus</span> type with <span style="font-style: italic;">Szechuanosaurus</span>
was followed by Molnar et al. (1990) where they consider the taxon an
allosaurid, which makes sense as Dong was a coauthor. Most recently,
Hendrickx et al. included <span style="font-style: italic;">Chienkosaurus</span>
in their cluster analyses, although the taxon is never mentioned in the
text, matrices or table of examined taxa. Classical/Hierarchical
clustering resolves it with <span style="font-style: italic;">Genyodectes</span>, <span style="font-style: italic;">Sinraptor dongi</span> (the only metriacanthosaurid analyzed there) and <span style="font-style: italic;">Allosaurus</span>, while neighbour joining clustering resolves it sister to a clade whose basal members are '<span style="font-style: italic;">Indosuchus</span>' AMNH jaws, <span style="font-style: italic;">Allosaurus</span> and <span style="font-style: italic;">S. dongi</span>. <br />
Young placed locality 47 at "the top part of the Kuangyuan Series and
immediately below the Chentsianyen conglomerate", now known as the
Guangyuan Group and the Chengqiangyan Group, with the former
corresponding to the Xiashaximiao Formation through the Penglaizhen
Formation. As it was found "immediately below" the boundary (layer 8b in Young et al., 1943), <span style="font-style: italic;">Chienkosaurus</span>
may be from the Penglaizhen Formation or slightly lower Shuining
Formation. The age is listed as Tithonian on fossilworks and in
Weishampel (1990), the latter cited as from "Dong (pers. comm.)". <br />
The tooth is similar to many large theropod teeth in general
characters, but is from the premaxilla as evidenced by the twisted
mesial carina and reduced extent of mesial serrations. The crown
base ratio (.75) is between the third and fourth premaxillary teeth of
<span style="font-style: italic;">Sinraptor dongi</span>'s holotype
(pm3 .60; pm4 1.04), which also match in size (pm3 FABL 16.87 mm; pm4
BW 12.26 mm) and in lacking mesial serrations basally. The cited
mesial (15 per 5 mm) and distal (6.7-10 per 5 mm) serration densities
are matched by teeth of <span style="font-style: italic;">S. dongi</span>,
and the strong mesial carina Young describes could easily be due to the
"longitudinal groove adjacent to the mesial carina, on the lingual
surface of the crown" "clearly present in lpm3 and lpm4" as described
by Hendrickx et al. (2020) for <span style="font-style: italic;">S. dongi</span>. Thus <span style="font-style: italic;">Chienkosaurus</span> is indistinguishable from <span style="font-style: italic;">Sinraptor dongi</span>
as far as can be determined from the description, and given its poorly
constrained age could be contemporaneous or even synonymous. Hendrickx
et al.'s matrices show no differences between <span style="font-style: italic;">Yangchuanosaurus shangyouensis</span> (including <span style="font-style: italic;">Y. magnus</span>), <span style="font-style: italic;">Sinraptor dongi</span> and <span style="font-style: italic;">S. hepingensis</span> that can be evaluated for <span style="font-style: italic;">Chienkosaurus</span>,
so pending Hendrickx's in prep. study on metriacanthosaurid dental
anatomy the genus is considered Metriacanthosauridae indet..<br />
<span style="font-weight: bold;">Referred material</span>- Young (1942) figured and described an ulna from the type locality (IVPP V193), stating he "would prefer to refer this ulna to <span style="font-style: italic;">Chienkosaurus ceratosauroides</span> above described." The element is very different from <span style="font-style: italic;">Limusaurus</span>
in having marked transverse expansions proximally and distally as well
as a triangular versus reniform proximal end, so that if <span style="font-style: italic;">Sinocoelurus</span> is closely related to that genus the ulna is unlikely to belong to it. <span style="font-style: italic;">Eoabelisaurus</span> has a much longer olecranon. The ulnae of megalosaurids and <span style="font-style: italic;">Kaijiangosaurus</span> is far more robust with more proximally extended olecranons, while those of most coelurosaurs (e.g. <span style="font-style: italic;">Zuolong</span>, <span style="font-style: italic;">Guanlong</span>, <span style="font-style: italic;">Coelurus</span>, <span style="font-style: italic;">Tanycolagreus</span>, <span style="font-style: italic;">Fukuivenator</span>) are much more slender with developed olecranons as well. <span style="font-style: italic;">Fukuiraptor</span>
has a dissimilar ulna with a strong olecranon, prominent anteroproximal
longitudinal ridge and unexpanded distal end. This leaves several
roughly comparable taxa whose ulnae have been figured in
anteroposterior view- <span style="font-style: italic;">Ceratosaurus</span>, <span style="font-style: italic;">Poekilopleuron</span>, <span style="font-style: italic;">Yangchuanosaurus</span>, <span style="font-style: italic;">Allosaurus</span> and <span style="font-style: italic;">Haplocheirus</span>. Young compared it favorably to the former, writing "it fits rather well with the ulna of <span style="font-style: italic;">Ceratosaurus nasicornis</span>
(length of ulna, 17.7 cm.) which is only slightly longer than the
present form", and indeed the main difference in profile is the more
gradual proximal expansion laterally. However, in proximal view IVPP
V193 differs from <span style="font-style: italic;">Ceratosaurus</span> and most other proximally figured ulnae in having a centrally placed olecranon (also seen in <span style="font-style: italic;">Coelurus</span>, but not <span style="font-style: italic;">Tanycolagreus</span>). While only photographed in anterior view, the ulna of <span style="font-style: italic;">Yangchuanosaurus</span> (CV 00214) would also seem to have a centrally placed olecranon, so IVPP V193 may be correctly referred to <span style="font-style: italic;">Chienkosaurus</span>/<span style="font-style: italic;">Szechuanosaurus</span> after all.<br />
Young
(1942) describes IVPP V190 as "A complete centrum of an anterior caudal
vertebra (or posterior lumbar) with length 66 mm., breadth 41 mm.,
minimum breadth of the centrum 24 mm", noting it "fit in size with <span style="font-style: italic;">Chienkosaurus ceratosauroides</span>" and calling it Theropoda indet. in the plate caption but also saying there it "probably belonging to <span style="font-style: italic;">Chienkosaurus ceratosauroides</span>." With a length/height ratio of 144% it is comparable to the ninth caudal of <span style="font-style: italic;">Sinraptor hepingensis</span>
and indistinguishable in lateral view. It differs in being 85% wider
than tall vs. 95%, but this is within the range of variation in <span style="font-style: italic;">hepingensis</span>'
caudals. Notably, this is from a different locality than the type,
said by Young to be in "the middle part of the" ... "Kuangyuan Series", layer 5a in Young et al. (1943),
and thus possibly corresponding to the Shangshaximiao Formation. Thus
while lacking a plausible connection to <span style="font-style: italic;">Chienkosaurus</span>, it is congruent with being metriacanthosaurid but may also be e.g. piatnitzkysaurid or megalosaurid.<br />
<b>References</b>- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan,
China. Bulletin of the Geological Society of China. 22(3-4), 293-309.<br />
Young, Bien and Mi, 1943. Some geologic problems of the Tsinling. Bulletin
of the Geological Society of China. 23(1-2), 15-34.<br />
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 1-772.<br />
Huene, 1959. Saurians in China and their relations. Vertebrata PalAsiatica. 3(3), 119-123.<br />
Steel, 1970. Part 14. Saurischia. Encyclopedia of Paleoherpetology. Gustav Fischer Verlag. 1-87.<br />
Rozhdestvensky, 1977. The study of dinosaurs in Asia. Journal of the Palaeontological Society of India. 20, 102-119.<br />
Dong, Zhang, Li and Zhou, 1978. [A new carnosaur discovered in
Yongchuan, Sichuan]. Chinese Science Bulletin. 23(5), 302-304.<br />
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.<br />
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 169-209.<br />
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. University of California Press. 63-139.<br />
Hendrickx, Stiegler, Currie, Han, Xu, Choiniere and Wu, 2020. Dental anatomy of the apex predator <span style="font-style: italic;">Sinraptor dongi</span> (Theropoda: Allosauroidea) from the Late Jurassic of China. Canadian Journal of Earth Sciences. 57(9), 1127-1147.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjh43UkPpd1Nb7HSMGpJzFHJJNkNSRSJ9f4fHRbL-rUUcZlfIvTpbbnMB0bJC7vSEXgEAu0GDJ3VjpFzifSKFjSp3Blz9tP0eNd8DKI1WhtBsotOwM5mNbK6HcAuR8-heZx88rUB6eeXKt6H2hzj45KMrDmrznOG6Bj7aQCXeQEpevtKGIzBlGWAJ6r/s447/Chienkosaurus.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="447" data-original-width="277" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjh43UkPpd1Nb7HSMGpJzFHJJNkNSRSJ9f4fHRbL-rUUcZlfIvTpbbnMB0bJC7vSEXgEAu0GDJ3VjpFzifSKFjSp3Blz9tP0eNd8DKI1WhtBsotOwM5mNbK6HcAuR8-heZx88rUB6eeXKt6H2hzj45KMrDmrznOG6Bj7aQCXeQEpevtKGIzBlGWAJ6r/s320/Chienkosaurus.png" width="198" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Lectotype tooth of <i>Chienkosaurus ceratosauroides</i> in mesial and basal views (after Young, 1942).<br /></td></tr></tbody></table><br /><p>Also newly revealed to be metriacanthosaurid is "Yuanmouraptor"-<br /><br /><b><a name="Yuanmouraptor"></a>"Yuanmouraptor"</b> Anonymous, 2014<br />
<b>Middle Jurassic<br />
Yuanmou County, Yunnan, China<br />
Material</b>- (ZLJ 0115) partial skull, mandibles (one incomplete, one partial),
postcrania<br />
<b>Comments</b>- This specimen is on display at the ZLJ as a new carnosaur,
but has yet to be described. There is a mounted skeleton, but how much is original
is unreported. Hendrickx et al. (2019) call this "an undescribed metriacanthosaurid (ZLJT 0115)", state it has mesial and
lateral teeth with "four to six, possibly more" flutes, and list it as
"Metriacanthosauridae indet." in their Appendix 1 indicating
information was from photos provided by Stiegler. <br />
<b>References</b>- Anonymous, 2014. Special Exhibition: Legends of the Giant
Dinosaurs. Hong Kong Science Museum newsletter. 1-3-2014, 2-7.<br />
Hendrickx, Mateus, Araújo and Choiniere, 2019. The distribution of
dental features in non-avian theropod dinosaurs: Taxonomic potential,
degree of homoplasy, and major evolutionary trends. Palaeontologia
Electronica. 22.3.74, 1-110.<br /><br /><br />As part of a review of Jurassic Chinese theropods, I went over the largest Jurassic theropod, previously referred to <i>Szechuanosaurus campi</i>.<br /><br /><span style="font-weight: bold;">unnamed averostran</span> <span style="font-style: italic;"><span style="font-weight: bold;"></span></span>(Camp, 1935)<br />
<b>Middle Jurassic?<br />Jung-Hsein UCMP V1501, middle Chongqing Group, Sichuan, China</b><br />
<b>Material</b>- (UCMP 32102) (~14.7 m) mesial dentary tooth (~69x~22x? mm), rib fragment, ischial fragment, femoral fragment (~1.33 m)<br />
<b>Comments</b>-
The specimen was collected on August 30 1915 by Louderback. Note the
UCMP locality number is V1501 (as determined in their online
catalogue), not V151 as listed by Camp. Jung-Hsien is now called
Rongxian, a county in Zigong City. Camp stated "The beds in which they
occur have been called the Szechuan series", which was a term for the
stratigraphic section from the Early Jurassic Qianfuyan (= Tsienfuyan)
Formation and Ziliujing (= Tsuliuching, = Tzeliutsin) Formation to the
Cretaceous Chengqiangyan (= Chengtsiangyen) Group and Jiading (=
Chiating, = Tshiating) Group, depending on north versus south in the
Sichuan Basin. Rongxian is located in the south, so UCMP V1501 would be
part of the Ziliujing-Chongqing-Jiading sequence, and Young (1937; see
also Young et al., 1943) placed it above the Ziliujing Formation but
below the conglomerates of the Jiading Group, and thus within the
Middle-Late Jurassic Chongqing Group. Furthermore, Young (1937) stated
"The fossiliferous horizon discovered by Louderback lies probably
between our horizons 2 and 3, some 200 meters above horizon 2" which is
the type locality of <span style="font-style: italic;">Omeisaurus junghsiensis</span>. As <span style="font-style: italic;">Omeisaurus</span>
is generally recovered in the Xiashaximiao Formation and horizon 3 is
another 300 meters above where Young placed UCMP V1501 (so may be the
Penglaizhen or Suining Formation), UCMP 32102 may derive from the
Shangshaximiao Formation. Dong et al. (1983) listed it as deriving
from that formation, perhaps using the same logic although they did not
describe any explanation.<br />
Camp (1935) initially referred the specimen to Megalosauridae because
histology "shows quite definitely that the relationship of the Chinese
form is with <span style="font-style: italic;">Allosaurus</span>" instead of <span style="font-style: italic;">Tyrannosaurus</span>. However, the plate shows this is because the sampled section of <span style="font-style: italic;">Tyrannosaurus</span> femur (labeled AMNH 5886, but this is the <span style="font-style: italic;">Anatotitan</span> paratype, and it is more probably <span style="font-style: italic;">Dynamosaurus</span> holotype AMNH 5866 that is known to be histologically sampled) is composed of secondary osteons, while those of <span style="font-style: italic;">Allosaurus</span> and UCMP 32102 are fibrolamellar bone. Yet <span style="font-style: italic;">Allosaurus</span>
can develop secondary osteons where bone is redeveloped as well (e.g.
MHNG GEPI V2567a), so this isn't a real difference between these taxa.
While Camp wrote "a projection of the borders would indicate an
original total length of at least 90 mm" for the tooth, he also
repeated Osborn's 1906 statement that <span style="font-style: italic;">Tyrannosaurus</span>
(CM 9380 and NHMUK R7994) teeth are up to 125 mm, which includes the
root. Combined with his statement the serrated distal carina "reaches
the base of the enamel" and serrations are not illustrated on the most
basal section, the actual crown length would have been about 69 mm.
Similarly, Camp wrote "At the base it is 17 mm. in longest diameter",
but scaling the figured tooth to the stated preserved length of 60 mm
results in a FABL of 22 mm instead. Young (1942) later wrote "The
general structure of [<span style="font-style: italic;">Szechuanosaurus campi</span>
syntype] V236 with the way of serrations fits so well with the
Junghsien tooth, we feel that there is practically no doubt in
regarding them as identical" "and prefer to consider the Junghsien
tooth as belonging also to the new form" <span style="font-style: italic;">Szechuanosaurus</span>. This despite previously stating UCMP 32102 "is bigger and straighter than all" <span style="font-style: italic;">Szechuanosaurus</span> syntype teeth. Compared to <span style="font-style: italic;">Sinraptor dongi</span> and <span style="font-style: italic;">Szechuanosaurus</span>,
UCMP 32102 is larger (~69 vs. up to 63 vs. ~32 and ~47 mm), with a much
greater crown height/base ratio (~314% vs. up to 244% vs. ~224% and
~267%), making it less tapered. The crown section is similar to <span style="font-style: italic;">Allosaurus</span>' fourth dentary tooth and the mid crown ratio of 53% is similar to dentary teeth in <span style="font-style: italic;">S. dongi</span> and between <span style="font-style: italic;">S. campi</span> IVPP V238B and 238C. As in mesial teeth of <span style="font-style: italic;">S. dongi</span>,
the crown is slightly lingually curved and the mesial carina does not
reach the crown base. The same serration densities (mesial 15 per 5
mm, distal 6.7-10 per 5 mm) can be found in <span style="font-style: italic;">S. dongi</span> as well, but are lower than <span style="font-style: italic;">S. campi</span>
(distal ~12-19 per 5 mm). Dong et al. (1983) wrote "Camp's description
and the dentition size suggests it may be assignable to <span style="font-style: italic;">Yangchuanosaurus</span>", but it is larger than even the <span style="font-style: italic;">magnus</span>
type (up to 75 mm), and more elongate than the largest maxillary teeth
of the genotype (crown height/base ratio of 267%), but detailed dental
statistics of the genus have yet to be published. The femoral fragment
is notably large, Camp stating the shaft has "an enormous hollow cavity
about 125 mm. in the longest diameter of its ellipse. The greatest
diameter of this segment at its narrowest point is 20 cm." Based on
the absence of a fourth trochanter or medial narrowing, the section is
just distal to the former structure. Here large theropod femoral
shafts are wider than deep, so 200 mm would be the width and the
figured depth is then ~143 mm. Scaling from the largest
metriacanthosaurid, <span style="font-style: italic;">Yangchuanosaurus magnus</span>,
results in a femoral length of ~1.33 meters, not far from Camp's
"estimated total length of about 140 cm." The ischium "consists of a
moderately hollow shaft spreading into a broader, solid plate", which
could describe most non-maniraptoran ischia, while the "tip of a large
rib" is not described. Notably, the ischium was found 37 meters from
the other material, so its association is less certain.<br />
Given the above information, UCMP 32102 is different from the <span style="font-style: italic;">Szechuanosaurus</span> syntypes and anteriorly straighter than <span style="font-style: italic;">Sinraptor</span>
as well, and is perhaps the largest known Jurassic theropod. As no
characters are outside the range of ceratosaurids, it is considered
Averostra incertae sedis here.<br />
<b>References</b>- Camp, 1935. Dinosaur remains from the province of Szechuan.
University of California Publications, Bulletin of the Department of Geological
Sciences. 23(14), 467-471.<br />
Louderback, 1935. The stratigraphic relations of the Jung Hsien fossil
dinosaur in Szechuan red beds of China. University of California
Publications. Bulletin of the Department of Geological Sciences.
23(14), 459-466.<br />
Young, 1937. New Triassic and Cretaceous reptiles in China (With some
remarks concerning the Cenozoic of China). Bulletin of the Geological
Society of China. 17(1), 109-120.<br />
Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan, China. Bulletin
of the Geological Society of China. 22(3-4), 293-309.<br />
Young, Bien and Mi, 1943. Some geologic problems of the Tsinling. Bulletin
of the Geological Society of China. 23(1-2), 15-34.<br />
Dong, Zhou and Zhang, 1983. Dinosaurs from the Jurassic of Sichuan. Palaeontologica
Sinica. Whole Number 162, New Series C, 23, 136 pp.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjOllwUZ06gVJCyS8N8hvzuSlUSLXT18-SZXUqYhABsYgOe9zf0f7wjDF9Sx5jDOkW8z2ugxvbcwJ6nd6yEMApD3EggnYTUBg-seFkDR4iN7DGZZUrlkYhlRjz_qytDGqisALVh8aVyDWp5g95rr879acMe9AuGO_7lL4hceB8a5IsF1O5VnwosfhGf/s1205/UCMP%2032102.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="941" data-original-width="1205" height="250" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjOllwUZ06gVJCyS8N8hvzuSlUSLXT18-SZXUqYhABsYgOe9zf0f7wjDF9Sx5jDOkW8z2ugxvbcwJ6nd6yEMApD3EggnYTUBg-seFkDR4iN7DGZZUrlkYhlRjz_qytDGqisALVh8aVyDWp5g95rr879acMe9AuGO_7lL4hceB8a5IsF1O5VnwosfhGf/w320-h250/UCMP%2032102.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">UCMP 32102 tooth (top) and femoral shaft (bottom) (after Camp, 1935).<br /></td></tr></tbody></table><p><br />Finally, I've been translating He (1984), so here's that publication's supposed <i>Szechuanosaurus</i>-<br /><br /><b>unnamed Tetanurae</b> (He, 1984)<br />
<b>Bathonian-Callovian, Middle Jurassic</b><br />
<b>Hexi Commune, Shangshaximiao Formation, Sichuan, China</b><br />
<b>Material</b>-
(CUT coll; = CCG coll) (multiple individuals) many teeth (~63 mm),
anterior cervical centrum (~68 mm; immature; Fig. 6-16, Pl. X Fig. 3),
mid cervical vertebra (~69 mm; Fig. 6-17), tenth cervical centrum
(immature; Fig. 6-19a), ~second dorsal centrum (~55 mm; immature?; Pl.
X Fig. 4), incomplete ~fourth dorsal vertebra (Fig. 6-19b), mid dorsal
centrum (immature; Fig. 6-19c), more than forty caudal vertebrae
including proximal caudal vertebra (Fig. 6-19d) and distal caudal
vertebra (~70 mm; Pl. X Fig. 5), incomplete coracoid (~98 mm
proximodistally), humerus (265 mm), ischium (~356 mm), femur, tibia
(~730 mm), fibula (~709 mm) and unguals<br />
<b>Comments</b>-
He (1984) states that in 1964, 1979 and 1980 in Chengdu the institute
(= CUT) "conducted systematic collections in Hexi Commune (near Huomu
Station) in the suburbs of Qingyuan City, including many carnosaur
specimens, including many teeth, cervical vertebrae, dorsal vertebrae,
more than forty caudal vertebrae, complete ischium, femur, tibia and
fibula, as well as relatively complete humerus, coracoid and claws."
(translated) He referred these to <span style="font-style: italic;">Szechuanosaurus campi</span>
because the syntypes were also found in the suburbs of Guangyuan and
believed to be from the Shangshaximiao Formation based on faunal
similarities and fossil abundance, "there is no significant difference
in shape and size" between <span style="font-style: italic;">S. campi</span>
and the Hexi teeth, and "there is no evidence of the existence of two
or more carnosaurs" from that horizon. However, the teeth of <span style="font-style: italic;">S. campi</span>have
not been shown to be diagnostic within e.g. Metriacanthosauridae,
multiple taxa with megalosaur-grade teeth are now known from the
Shangshaximiao (<span style="font-style: italic;">Leshansaurus</span>, <span style="font-style: italic;">Yangchuanosaurus shangyouensis</span>, <span style="font-style: italic;">Sinraptor hepingensis</span>), and <span style="font-style: italic;">S. campi</span> itself may be from the Penglaizhen Formation or slightly lower Shuining
Formation instead. Furthermore, He notes "that the tooth size in this batch of <span style="font-style: italic;">Szechuanosaurus campi</span>
material we collected is quite varied, which means that in addition to
the differences in individual size, there may also be immature
specimens, because some vertebral centra and neural arches are unfused.
The largest individual is comparable to the type of <span style="font-style: italic;">Yanchuanosaurus shangyouensis</span>
[sic], and the smallest individual is estimated to be only 4-5 meters
in length." Thus multiple individuals and perhaps multiple taxa
are involved, with only the tibia and fibula in Plate X Figures 8-10
being claimed to be from one individual. Note while Chure (2000)
mentioned a metatarsal as being in this material, He does not indicate
as such and Chure might have mistaken Plate X Figure 6 which is a
humerus. Indeed, Chure seems not to have translated the text so
understates the preserved vertebral number and misses the reference to
unguals. Yang et al. (2021) later describe the humeral histology, noting their <span style="font-style: italic;">Szechuanosaurus</span> specimen is from Hexi and citing He's paper. This paper confirms the material "<span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">contains several incomplete individuals with large differences in size" (translated), that "</span></span></span><span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">The
specimen is currently preserved in the Museum of Chengdu University of
Technology" and that it was recovered from the Shangshaximiao Formation
at the same locality as <span style="font-style: italic;">Mamenchisaurus</span> "guangyuanensis". "</span></span></span><span class="HwtZe" lang="en"><span class="jCAhz ChMk0b"><span class="ryNqvb">One
of the medium-sized individuals was recovered, mounted and exhibited",
which is photographed in their Figure 2, although it cannot be
determined what material is real and what is plaster.</span></span></span><br />
This material was originally referred to Megalosauridae by He (1984)
based on tetanurine plesiomorphies (large teeth, short presacral
vertebrae, distally expanded ischium), while Chure (2000) placed it in
non-avetheropod Tetanurae based on the supposed lack of a posterodistal
coracoid process and subglenoid fossa, although Figure 6-18 of He
clearly shows both. Although Chure believes the information
available in the literature "make(s) it impossible to refer this
material to any family" and considered it indeterminate, the figures
and plates suggest otherwise. Among Late Jurassic theropods, the
slightly opisthocoelous cervicals are only known in piatnitzkysaurids
and coelurosaurs, with the long and low neural spines being unlike most
contemporary non-coelurosaur theropods, meaning the mid cervical
vertebra at least is not megalosauroid or carnosaurian.
Similarly, the large coracoid tubercle is unlike basal tetanurines and
more similar to ceratosaurs or coelurosaurs, although lacking the
hypertrophied size of the former. As suggested by Chure, the
ischium does resemble <span style="font-style: italic;">Megalosaurus</span> in the ventral kink of the shaft and boot morphology, although it is much more robust, thus a referral to <span style="font-style: italic;">Leshansaurus</span> is plausible. The tibia on the other hand is more similar to <span style="font-style: italic;">Sinraptor</span>
in the anteroposteriorly short proximal end and anteroposterior
compression distally, so may be metriacanthosaurid. Based on this
brief comparison, the material deserves restudy and probably represents
multiple tetanurine taxa.<br />
<b>References</b>- He, 1984. The Vertebrate Fossils of Sichuan. Sichuan Scientific
and Technical Publishing House, Chengdu, Sichuan. 168 pp.<br />
Chure, 2000. A new species of <i>Allosaurus</i> from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. PhD thesis. Columbia University. 964 pp.<br />
Yang, Liu and Zhang, 2021. The humeral diapophyseal histology and its biometric significance of Jurassic <span style="font-style: italic;">Szechuanosaurus campi</span> (Theropoda, Megalosauridae) in Guangyuan City, Sichuan Province. Acta Geologica Sinica. 95(8), 2318-2332.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqkhCklSgPC2PPGweRNYdbOTLygaOcadwdPDR8Kyj5A8zA_LHt05ScYrAGDOAohABt3EpFSYBs6ksx0ZnXC6gnkW403CkX_bYDsrnmtZbQdH2nkliOH1ZpgPKjpDcNI8FuuHB_1iiHzS1SceN4IyCRPuF2Wp54ogIrAW_IUAP7RMBAd0pyLuNYfJqi/s1004/He%201984%20Szechuanosaurus.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1004" data-original-width="660" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqkhCklSgPC2PPGweRNYdbOTLygaOcadwdPDR8Kyj5A8zA_LHt05ScYrAGDOAohABt3EpFSYBs6ksx0ZnXC6gnkW403CkX_bYDsrnmtZbQdH2nkliOH1ZpgPKjpDcNI8FuuHB_1iiHzS1SceN4IyCRPuF2Wp54ogIrAW_IUAP7RMBAd0pyLuNYfJqi/s320/He%201984%20Szechuanosaurus.png" width="210" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Tetanurae elements from He (1984)- 1, 2 teeth; 3 anterior cervical centrum; 4 ~second dorsal centrum; 5 distal caudal vertebra; 6 humerus; 7 ischium; 8 tibia in anterior and distal views; 9 fibula in lateral and distal views; 10 tibia and fibula in proximal view (after He, 1984).<br /></td></tr></tbody></table><p><br /><b>Additional references</b>- Siegwarth, Lindbeck, Redman, Southwell, unpublished. Giant carnivorous dinosaurs
of the family Megalosauridae from the Late Jurassic Morrison Formation of eastern
Wyoming. Contributions from the Tate Museum Collections, Casper, Wyoming. 2,
40 pp.<br /><br />Benson, 2010b. The osteology of <i>Magnosaurus nethercombensis</i> (Dinosauria,
Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination
of the oldest records of tetanurans. Journal of Systematic Palaeontology. 8(1),
131-146. <br /><br />Benson, 2010b (online 2009). A description of <i>Megalosaurus bucklandii</i> (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.</p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com11tag:blogger.com,1999:blog-3248412803814730250.post-68047089960935219162022-07-05T20:34:00.000-07:002022-07-05T20:34:01.180-07:00The Theropod Database July update - Iren Dabasu theropods<p>As mentioned by Justin Tweet last month, we haven't had that many new dinosaur taxa described lately, whether due to COVID or random chance. But this doesn't bother me because there is always so much data to catch up on. In mid May I checked my Database suggestion emails and the most recent topic to address was "Brochu (2003) mentions that the specimen AMNH 6266 (including a jugal,
lacrimal, quadratojugal, and D-shaped premaxillary tooth) might belong
to the holotype of <i>Alectrosaurus olseni </i>as it was found at the type locality of <i>A. olseni</i>." And because of that, we now have all published Iren Dabasu theropod records added and updated on The Theropod Database. I'll go over some of the lesser known and new details about select taxa here. There was nothing particularly new about <i>Erliansaurus</i>, <i>Neimongosaurus </i>or <i>Gigantoraptor</i>, so I don't list them below. If anyone knows of any Iren Dabasu theropod specimens in the literature I missed, do leave a comment. One thing that struck me was just how much material remains undescribed, at the AMNH, IVPP and PIN especially. If anyone needs projects based on collected material- sort out <i>Archaeornithomimus</i>, describe the many therizinosaur elements at the IVPP, describe the partial troodontid skeletons at the AMNH, etc..<br /></p><p><a href="https://www.theropoddatabase.com/Tyrannosauroidea.html#Alectrosaurusolseni"><b><i>Alectrosaurus olseni</i></b></a><br /><b>(see page for materials list)</b><br />The lectotype hindlimb AMNH 6554) was discovered on April 25 1923 at
Third Asiatic Expedition field site 136, while a partial forelimb (AMNH
6368) was found on May 4 at field site 138, 30 meters away. Andrews
(1932) first mention the former as "the complete hindlimb of a large
carnivorous dinosaur. The leg lay doubled up just as the great reptile
had died millions of years ago." Gilmore (1933) made each a syntype of
his new taxon of deinodontid, <span style="font-style: italic;">Alectrosaurus olseni</span>,
noting "in the field they were thought to pertain to the same
individual." but that he preferred to treat them as two individuals.
They were considered the same taxon based on the manual unguals
(questionably associated in the case of AMNH 6554) "being laterally
compressed, strongly curved, and having sharply pointed extremities",
which are characters broadly true of almost all theropod manual
unguals. Barsbold (1976) was the first to consider AMNH 6368 wrongly
assigned, stating "As new materials from the MPR* show, a large ungual
phalanx previously attributed to the manus of <span style="font-style: italic;">Alectrosaurus</span>
(Gilmore, 1933) does not really belong to it" (translated), citing the
still undescribed IGM 100/50 from Bayanshiree which includes "a small
ungual phalanx of the first manual digit, quite typical for
tyrannosaurids." Further, he noted "A large, laterally compressed
ungual phalanx, similar in structure and form to that attributed to <span style="font-style: italic;">Alectrosaurus</span>,
belongs to another previously unknown dinosaur (under study) found
there. This dinosaur does not belong to Tyrannosauridae.", which is a
reference to the also undescribed <span style="font-style: italic;">Segnosaurus</span>.
In their redescription, Mader and Bradley (1989) describe AMNH 6368 in
detail and place it in Segnosauridae, and it has been viewed as
therizinosaurian since then. As explained by Mader and Bradley, the
type listing by White (1973) combined the syntype materials, so that
Welles and Long (1974) officially declared the hindlimb as the
lectotype when they stated "we here designate this specimen, AMNH 6554,
the type of the species." Zanno (2010: Fig. 9D) figured the manual
unguals of AMNH 6554 as therizinosaurian without comment, although
their non-tyrannosaurid characters could also be plesiomorphically
shared with e.g. <span style="font-style: italic;">Dryptosaurus</span>.<br />Dong et al. (1989) first reported <span style="font-style: italic;">Aublysodon</span>
from the July 1988 Sino-Canadian expedition (CCDP), and Currie et al.
(1990) stated "Identical teeth [to Dinosaur Park juvenile tyrannosaurid
'<span style="font-style: italic;">Aublysodon</span>'] recently were recovered from the Iren Dabasu Formation at Erenhot, People's Republic of China (IVPP 170788104). The Asian "<span style="font-style: italic;">Aublysodon</span>" teeth belong to <span style="font-style: italic;">Alectrosaurus</span>
(Perle pers. comm. 1989..." Similarly, Dong (1992) reports "In July
1988, the expedition of the CCDP came to Erenhot (Fig.85) where they
collected ... teeth of ... large theropods (tyrannosaurid)." Dong et
al. first reported that in July 1988 "A partial skeleton of <span style="font-style: italic;">Alectrosaurus</span> was discovered too late to collect", and Dong (1993) followed that up by writing "An incomplete skeleton of <span style="font-style: italic;">Alectrosaurus</span>
was found by Currie [in 1988], but was not excavated until the return
expedition of 1990", which was at CCDP site #9 based on Currie and
Eberth's (1993) table 3. They further noted "Perle (1977) ... has been
studying more recently discovered postcranial specimens (Perle, pers.
comm. 1989)" and that "The absence of denticles on the premaxillary
teeth (Perle, pers. comm. 1989; IVPP 180788-104) suggests that it
should be included in the Aublysodontinae." The similar field numbers
to Currie et al.'s suggest one is a typo, and serrationless
premaxillary teeth are a juvenile character of tyrannosaurines but also
known in some basal tyrannosauroids (<span style="font-style: italic;">Yutyrannus</span>, <span style="font-style: italic;">Xiongguanlong</span>). Currie (2001) reported "Several partial, undescribed skeletons of <span style="font-style: italic;">Alectrosaurus</span>
collected from southeastern Mongolia are in the collections of the
museum in Ulaanbaatar, and another new specimen was recently collected
from Erenhot in China", the latter seemingly being the one mentioned by
Dong et al..<br /><b>(see page for references)</b><br /></p><p><span style="font-weight: bold;">undescribed tyrannosauroid</span> (Granger and Berkey, 1922)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material</b>-
(AMNH 6266; "AMNH 6556" of Carr, 2005) (small) incomplete lacrimal,
anterior jugal, quadratojugal, lateral ectopterygoid, posterior
pterygoid, premaxillary teeth, lateral teeth<br />
<span style="font-weight: bold;">Comments</span>- Brochu (2003) noted "A box of bone fragments (AMNH 6266) from the same locality [as <span style="font-style: italic;">Alectrosaurus</span>]
includes small tyrannosaurid skull bones (including a characteristic
jugal, lacrymal, quadratojugal, and D-shaped premaxillary tooth) that
might belong to the same individual. ... The skull parts are consistent
with <span style="font-style: italic;">Albertosaurus</span>; for example, the jugal foramen is a dorsally-opening slit." He states it "had been originally catalogued as "<span style="font-style: italic;">Deinodon</span>
sp." ..., but this was subsequently scratched off and "Theropoda
indet." written on in pencil", and the AMNH online catalogue does list
6266 as "<span style="font-style: italic;">Deinodon</span>
? sp." Carr (2005) later reported "an undescribed, but shattered,
tyrannosauroid skull (AMNH 6556) from the same general area - Iren
Dabasu -" as the <span style="font-style: italic;">Alectrosaurus</span>
lectotype, but as they are from different locations "there is no
evidence they are from the same individual." Carr states "The presence
of a secondary fossa in the antorbital fossa of the jugal indicates the
specimen is referable to Tyrannosauridae. The cornual process of the
lacrimal is similar to some juvenile tyrannosaurids in that it is a
low, laterally extending ridge. The lateral teeth are as finely
denticulate as tyrannosauroid teeth of the same basal crown length from
the Turonian of Uzbekistan." This is the "Iren Dabasu taxon" in Carr's
tyrannosauroid analyses as of Carr et al. (2017), under study by Carr
and recovered in a
polytomy with <span style="font-style: italic;">Timurlengia</span>, <span style="font-style: italic;">Jinbeisaurus</span>
and eutyrannosaurs as of 2019. While this is certainly the same
specimen based on material preserved, the AMNH online catalog lists
this specimen number as being a saurischian metatarsal II with a
locality "8 mi. E. of station" which would place it among Third Asiatic
Expedition field sites 140-149 (while <span style="font-style: italic;">Alectrosaurus</span>
is from 136 less than a mile south of the station). AMNH 6556 is
listed in the catalogue as collected on April 30, which matches Carr's
statement the skull was found in late April five days apart from <span style="font-style: italic;">Alectrosaurus</span>'
lectotype (which was found on April 25). However, Mehling (pers. comm.
6-2022) indicates AMNH 6556 is actually a metatarsal II and that AMNH
6266 was discovered in 1922, so that Carr apparently got the specimen
number wrong and incorrectly used the metatarsal's discovery date for
the skull. The early discovery makes sense considering the low
specimen number and allows us to equate the material with "portions of
a small carnivorous dinosaur skull with two or three teeth" found in
the 1922 expedition as reported by Granger and Berkey (1922) along with
ornithomimid remains that are near certainly AMNH 6267-6268. If it was
recovered with the latter specimens, AMNH 6266 would have been found
between April 25 and May 7 at one of the western AMNH quarries
(131-138), and thus may be from the same locality the <span style="font-style: italic;">Alectrosaurus</span>
type as stated by Brochu. Alas, the only recorded locality information
in the AMNH card catalogue is Iren Dabasu (Mehling, pers. comm. 6-2022).<br />
<b>References</b>-
Granger and Berkey, 1922. Discovery of Cretaceous and older Tertiary strata in Mongolia. American Museum Novitates. 42, 7 pp.<br />
Brochu, 2003. Osteology of <i>Tyrannosaurus rex</i>: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
Society of Vertebrate Paleontology Memior. 7, 138 pp. <br />
Carr, 2005. A reappraisal of tyrannosauroids from Iren Dabasu, Inner Mongolia,
People's Republic of China. Journal of Vertebrate Paleontology. 25(3), 42A.<br />
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.</p><p><b>undescribed Tyrannosauridae</b> (Gilmore, 1933)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material</b>- (AMNH 21552) (large) femur (Mader and Bradley, 1989)<br />
(AMNH coll.) (large) pedal elements (Gilmore 1933)<br />
(IVPP coll.) teeth and/or elements (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)<br />
<b>Comments</b>- Under Deinodontidae, Gilmore (1933) states "The presence of a second carnivore, apparently rivaling <span style="font-style: italic;">Tyrannosaurus</span>
in size, is indicated by a few scattered foot bones." These are no
doubt one or more of the specimens listed here under undescribed
Averostra (e.g. AMNH 6376, 6556, 6744, 6756, 6757, etc.).<br />
Mader and Bradley (1989) noted "among the materials brought back by the
Central Asiatic Expeditions was the isolated femur (AMNH 21552) of a
much larger tyrannosaur [than the <span style="font-style: italic;">Alectrosaurus</span> lectotype]." This may belong to the same taxon as Gilmore's pedal material based on size.<br />
Yao et al. (2015) note "small unarticulated bones and teeth, including
fossils of ... tyrannosauroids" from "a rare microvertebrate locality
within the Iren Dabasu Formation, about 16 km northeast of Erenhot
City."<br />
<b>References</b>- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.<br />
Mader and Bradley, 1989. A redescription and revised diagnosis of the syntypes
of the Mongolian tyrannosaur <i>Alectrosaurus olseni</i>. Journal of Vertebrate
Paleontology. 9(1), 1-55.<br />
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. <i>Caenagnathasia</i>
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj8iIGvZQkYSIevsjdA_NLfZ6bopBsEfj8JYEo4WJVZxbw27MXnlBLDzEfG6Q3-zgrx7sYDy0iXtmjRZGJPzYO05uS31vCppChzpztc5fA8ykc_bTDJIhNkop9F1LxKstuyZ_2RWxSnC2iz2Rd9BgP8erp-1gbKwWMBWCNjvTHv3FLYuAaVFCuHyPiR/s3164/Archaeornithomimus%20pubes.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1030" data-original-width="3164" height="104" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj8iIGvZQkYSIevsjdA_NLfZ6bopBsEfj8JYEo4WJVZxbw27MXnlBLDzEfG6Q3-zgrx7sYDy0iXtmjRZGJPzYO05uS31vCppChzpztc5fA8ykc_bTDJIhNkop9F1LxKstuyZ_2RWxSnC2iz2Rd9BgP8erp-1gbKwWMBWCNjvTHv3FLYuAaVFCuHyPiR/s320/Archaeornithomimus%20pubes.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Iren Dabasu ornithomimosaur pubes (left to right) - AMNH 21799 figured by Smith and Galton (1990) as <i>Archaeornithomimus asiaticus</i> so generally associated with the taxon; AMNH 21798 and a distal pubis in AMNH 6570, both among <i>Archaeornithomimus </i>material from the same bonebed as 21799; LH-02-01 from an unknown locality in the same area described by Yao et al. (2022) as Ornithomimosauria indet.. The first three are my photos courtesy of the AMNH (scale = 100 mm), the fourth is after Yao et al., 2022.<br /></td></tr></tbody></table><br /></p><p><a href="https://www.theropoddatabase.com/Ornithomimosauria.htm#Archaeornithomimusasiaticus"><i><b>Archaeornithomimus asiaticus</b></i></a><br /><b>(see page for materials list and additional comments)<br />Comments</b>- Watanabe et al. (2015) state "Based on manual articulation of
disarticulated sacral vertebrae, we identify the two sacral vertebrae
(AMNH FARB 21790) sampled for this study as the second and third sacral
vertebrae", but these are the second and third caudals as identified by
Smith and Galton as confirmed by personal observation when sacrals 2-5
were on loan articulated to the ilium. Similarly, two of the proximal
caudals Watanabe et al. call AMNH 21790 are actually the last two
vertebrae in AMNH 21802.<br /> <span style="font-weight: bold;">Other material</span>- The AMNH catalog lists 6267 and 6268 as <i>Ornithomimus sp.</i>, but they are
probably <i>Archaeornithomimus</i>
based on stratigraphy. Given their lower specimen numbers, it is
likely these were recovered in the initial reconnaissance expedition to
Iren Dabasu in 1922 (April 25 to May 7) and represent the "Carnivorous
dinosaurs of at least two genera, the smaller one being of the <span style="font-style: italic;">Ornithomimus</span> type" reported by Granger and Berkey (1922). Indeed, those authors later state "Remains of the small <span style="font-style: italic;">Ornithomimus</span>-like
creature are particularly abundant and the last day at Iren Dabasu we
picked up probably fifty good foot bones and centra from two or three
knolls", which matches the listed material. The locality of AMNH 6267
is listed as "Iren Dabasu Sta. 1 1/2 SW of auto trail", so was
plausibly from one of the western AMNH quarries (131-138), as opposed
to the type material that was from the Kaisen Quarry (AMNH locality
140) and Johnson Quarry (AMNH locality 141) which were discovered the
following year. The AMNH online catalogue also lists AMNH 21626 and
21627 as possibly referrable to <i>Archaeornithomimus</i>. AMNH
21597 is figured as therizinosaurid ungual on the AMNH online catalog,
but the low curvature and distally placed flexor tubercle are instead
almost identical to <span style="font-style: italic;">Archaeornithomimus</span>
(e.g. AMNH 6570, 6576). Currie and Eberth (1993) state "The present
whereabouts of a partial skull found by the Sino-Soviet expedition is
currently unknown", but their paper's details indicate it was found in
June 1959 from their localities K (= AMNH locality 141?), L or P, and
initially stored in either the IVPP or PIN. Chow and Rozhdestvensky
(1960) specified the timing of excavations to be June 14 to July 17 and
state "materials collected include ... small ornithopods (of <span style="font-style: italic;">Struthiomimus</span>
type)", with 'ornithopods' presumably a typo for 'theropods', and
Currie and Eberth state "more than a thousand bones" were identified as
ornithomimid in the filed from the Sino-Soviet expedition. Currie and
Eberth also say that after the joint BMNH - Inner Mongolian Museum
expeditions of 1972-1977, "Some of the specimens (including
ornithimimid ... skeletons) were prepared for display in Hohhot", with
casts at the Erenhot Dinosaur Museum. These were mostly from
localities on the west side of Iren Nor. Dong (1992) noted that "<span style="font-style: italic;">Archaeornithomimus</span>
is the most common species in the bone beds" where the CCDP excavated
in July 1988 (and later in 1990), which would have ended up in the IVPP
as no theropod body fossils from Iren Dabasu are at the TMP. A distal
caudal described by Makovicky (1995) as <span style="font-style: italic;">Avimimus</span> "has the morphology of a typical
coelurosaurian distal caudal, but is otherwise undiagnostic. The
possibility that it may originate from an avimimid is suggested by its
small size. It should be noted, however, that it could just as
conceivably be from the tail of a juvenile <span style="font-style: italic;">Archaeornithomimus</span> from the same bonebed." Indeed, the <span style="font-style: italic;">A. nemegtensis</span> bonebed show <span style="font-style: italic;">Avimimus</span> has short distal caudals like other caenagnathoids and unlike ornithomimosaurs, so this specimen is here referred to <span style="font-style: italic;">Archaeornithomimus</span>. Godefroit et al. (1998) reported "one single ornithomimid caudal vertebra" from the <span style="font-style: italic;">Bactrosaurus</span>
bonebed they described from Locality SBDE 95E5, slightly to the west of
AMNH locality 140. Yao et al. (2015) note "small unarticulated bones
and teeth, including
fossils of ... ornithomimids" from "a rare microvertebrate locality
within the Iren Dabasu Formation, about 16 km northeast of Erenhot
City."<br /><span style="font-weight: bold;">More than one taxon?</span> There is non-ornithomimid material catalogued under <i>Archaeornithomimus</i>,
including a juvenile ?<span style="font-style: italic;">Bactrosaurus</span> ungual (in AMNH 6576),
a small ?troodontid pedal ungual I (in AMNH
6576) and part of an ?oviraptorid manual ungual I (in AMNH 6570). <br />
Yao et al. (2022) described pelvis and sacrum LH-02-01 discovered in
2002 from an undocumented locality near Iren Nor. They added the
specimen to Choiniere's coelurosaur analysis to recover it as an
ornithomimosaur in a polytomy with <span style="font-style: italic;">Nqwebasaurus</span>, <span style="font-style: italic;">Pelecanimimus</span>, <span style="font-style: italic;">Shenzhousaurus</span>, <span style="font-style: italic;">Beishanlong</span> and Deinocheiridae+Ornithomimidae. Supposed differences from <span style="font-style: italic;">Archaeornithomimus</span>
are- larger size (ilium 342 vs. 114 mm); "the shortest sacral centrum
is the second rather than the third [actually the third instead of the
fourth, assuming six sacrals as in <span style="font-style: italic;">Archaeornithomimus</span>-
Makovicky, 1995]; the centrum of the first caudal is shorter than that
of the fifth [sixth] sacral"; first caudal with flat posterior
articular surface vs. concave; first caudal with flat ventral surface
vs. median groove; first caudal neural spine posteriorly sloped vs.
vertical; pubic shaft straight vs. strongly posteriorly curved;
transition between anterior pubic shaft and dorsal edge of pubic foot
rounded vs. angled; anterior pubic foot more pointed; obturator process
less prominent; ischial foot limited to anterior expansion vs. expanded
some posteriorly; ischiopubic ratio 82% vs. 92%. However, it seems the
authors depended on the literature for their information on <span style="font-style: italic;">Archaeornithomimus</span>
(e.g. thinking the ischia are AMNH 21798 instead of 6558, and that the
first caudal would be amphicoelous and grooved because Smith and Galton
said in the proximal caudal paragraph "The centra are amphicoelous ...
and each has a shallow ventral groove"), and this led to misinformation
and incomplete statements. Regarding size, there is an ilium ~380 mm
long in AMNH 6570, larger than even LH-02-01. The proximal five
caudals of AMNH 21790 all lack median grooves ventrally and the
posterior articular surface of the first centrum is still attached to a
broken off part of the second centrum so cannot be evaluated. The
first caudal neural spine has been broken away since Smith and Galton's
drawing, and the shape of the other neural spines differ from the
drawing in greater to lesser degrees. There is a distal pubis in AMNH
6570 with an almost identical foot to LH-02-01, including the rounded
anterodorsal edge, more pointed anterior foot and shorter posterior
foot. Finally, there is no indication pubes AMNH 21799 and ischia AMNH
6558 were associated, and indeed five right proximal tibiae are known
from site 140 that can differ in size ~10% from each other. Thus sacra
AMNH 21790 and LH-02-01 may be distinct based on central lengths, pubes
AMNH 21799 and LH-02-01 / AMNH 6570 (in part) seem to be distinct, as
do ischia AMNH 6558 and LH-02-01. However, undescribed pubes AMNH
21798 are intermediate in having a slight anterior bow, rounded
anterodorsal transition, less pointed anterior foot and intermediate
posterior foot length. This suggests the sacral and ischial
differences could easily be due to the low sample size (N = 2) of each
element and that considering all of these specimens to be conspecific
is the most realistic and functional conclusion. Since the lectotype
is a partial pes not comparable to LH-02-01 at all (and without any
proposed diagnosis or comparison to other Iren Dabasu ornithomimosaur
pedal specimens), the other rational option would be to limit <span style="font-style: italic;">Archaeornithomimus asiaticus</span>
to AMNH 6565 and potentially other comparable specimens (e.g. AMNH
6568, 21616, 30240B/C and parts of 6570 and 6576) while leaving the
non-pedal specimens as Ornithomimosauria indet.. Alternatively,
LH-02-01 and e.g. AMNH 21790 or 6558 could be given names, but again
the vast majority of specimens would be incomparable and relegated to
Ornithomimosauria indet.. Given this data and pending the description
of some of the multitude of unpublished specimens, all Iren Dabasu
ornithomimosaurs are listed here under <span style="font-style: italic;">Archaeornithomimus asiaticus</span>.<br /><b>(see page for references)</b></p><p><b>unnamed therizinosaurid</b> (Gilmore, 1933)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<span style="font-weight: bold;">AMNH 138,</span><b>
Iren Dabasu Formation, Inner Mongolia, China</b><br />
<b>Material</b>- (AMNH 6368; syntype of <i>Alectrosaurus olseni</i>) humerus
(390 mm), manual ungual I (190 mm on curve), manual phalanx II-1 (74 mm)<br />
?...
(AMNH 21784) three mid caudal vertebrae (~41, ~38, ~35 mm), distal caudal centrum (~24 mm) (Mader and Bradley, 1989)<br />
?... (uncollected) two or three elements (Mader and Bradley, 1989)<br />
<b>Comments</b>- AMNH 6368 was found on May 4 1923 at Third Asiatic Expedition field site 138, 30 meters away from the lectotype of <i>Alectrosaurus
olseni</i> and was thought to belong to the same individual in the field.
Gilmore (1933) prefered to treat them as two individuals. He placed them in
the same species due to the similarity between the manual unguals, which are
“laterally compressed, strongly curved and have sharply curved extremities”,
as well as the association in the field and slenderness. Gilmore originally diagnosed
<i>A. olseni</i> partially on the characteristics of this specimen. Rozhdestvensky (1970) recognized the similarity to <span style="font-style: italic;">Therizinosaurus</span> and referred <span style="font-style: italic;">Alectrosaurus</span> to Therizinosauridae, but Barsbold (1976) and Perle (1977) correctly removed the forelimb from that taxon based on supposed <span style="font-style: italic;">Alectrosaurus</span>
specimen IGM 100/50 with small humerus and manual ungual. Mader
and Bradley (1989) described the specimen in detail as a segnosaurid.
Zanno (2010) notes the slender proportions, poorly defined medial
tuberosity, and posterior humeral trochanter indicates it is not
referable to <i>Neimongosaurus</i>, and it is more gracile than <i>Erliansaurus</i> and lacks
that genus' distinctive crest-shaped posterior trochanter. Zanno (2006) recovered it as more derived than <span style="font-style: italic;">Alxasaurus</span> in her phylogenetic analysis, and most recently Hartman et al. (2019) found it sister to <span style="font-style: italic;">Segnosaurus</span> in Therizinosauridae.<br />
AMNH 21784 was also discovered on May 4 1923 at Third Asiatic Expedition
field site 138, but
not catalogued until 1984 and not described until 1989 by Mader and
Bradley. While those authors described them as Theropoda incertae
sedis, they may be therizinosauroid based on their resemblence to <span style="font-style: italic;">Alxasaurus</span>'
mid caudals (e.g. short centra that become shorter distally; low neural
spines; similarly placed transverse processes) although their neural
spines are more posteriorly restricted and their prezygapophyses are
longer. Although Mader and Bradley considered the caudals too small to
belong to the same individual as the humerus, they were comparing them
to theropod proximal caudals as they were unaware of therizinosauroids'
modified caudal series with relatively homogenous caudals decreasing in
length distally. In fact, given that <span style="font-style: italic;">Alxasaurus</span>'
holotype humerus is 4% longer than AMNH 6368, its fourteenth-eighteenth
caudals are also slightly longer (39-44 mm) as is an isolated distal
caudal (26 mm), so the sizes are appropriate to belong to a single
therizinosauroid individual. As they are from the same field site,
that is provisionally accepted here. Mader and Bradley also noted "two
or three unspecified [theropod] elements that regrettably were not
collected due to their "poor condition" (Granger's field record, Third
Asiatic Expedition)" found with the forelimb and caudal material and
might also belong to the same individual.<br />
<b>References</b>- Gilmore, 1933. On the Dinosaurian Fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.<br />
Rozhdestvensky, 1970. Giant claws of enigmatic Mesozoic reptiles. Paleontological
Journal. 1970(1), 131-141. <br />
Barsbold, 1976. New data on <i>Therizinosaurus</i> (Therizinosauridae, Theropoda). In Kramarenko, Luvsandansan, Voronin, Barsbold, Rozhdestvensky, Trofimov and Reshetov (Eds.).
Paleontology and Biostratigraphy of Mongolia. The Joint Soviet-Mongolian
Paleontological Expedition, Transactions. 3, 76-92.<br />
Perle, 1977. On the first discovery of <i>Alectrosaurus</i>
(Tyrannosauridae, Theropoda) from the Late Cretaceous of Mongolia. Problemy
Geologii Mongolii. 3, 104-113.<br />
Mader and Bradley, 1989. A redescription and revised diagnosis of the syntypes
of the Mongolian tyrannosaur <i>Alectrosaurus olseni</i>. Journal of Vertebrate
Paleontology. 9(1), 41-55.<br />
Zanno, 2006. The pectoral girle and forelimb of the primitive therizinosauroid
<i>Falcarius utahensis</i> (Theropoda, Maniraptora): Analyzing evolutionary
trends within Therizinosauroidea. Journal of Vertebrate Paleontology. 26(3),
636-650. <br />
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of
Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic
Palaeontology. 8(4), 503-543.<br />
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. <br /></p><p><b>undescribed Therizinosauroidea</b> (Dong, 1992)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China</b><br />
<b>Material</b>- (Erenhot Dinosaur Museum coll.) dentary, teeth (Currie and Eberth, 1993)<br />(IVPP coll.) (isolated) many limb elements (Dong, 1992)<br />
(IVPP and PIN coll.) tens of specimens (Cuirrie and Eberth, 1993)<br />
<span style="font-weight: bold;">Comments</span>- Currie and Eberth
(1993) stated "A rough tally of Sino-Soviet field
identifications shows that ... 'theropods' (including large theropods,
small theropods and segnosaurs, but not ornithomimids) were more common
(400 specimens)" and that "The apparent high numbers of carnivorous
dinosaurs can be attributed mostly to ornithomimids [>1000 elements]
and segnosaurids" indicating some significant number of those 400
'theropods' found in June-July 1959 were therizinosaurs.<br />
Dong (1992) reported "In July 1988, the expedition of the CCDP came to
Erenhot (Fig.85) where they collected ... Many limb bones [which] might
be identified as segnosaurs" and listed <span style="font-style: italic;">Segnosaurus</span>
sp. as being present in the formation. He later (1993) said "at least
two taxa of segnosaurs" were represented. Currie and Eberth (1993)
stated "isolated elements were commonly recovered by the Sino-Canadian
expeditions in 1988 and 1990. A well-preserved dentary with teeth is in
the collections of the Erenhot Dinosaur Museum. The isolated elements
are indistinguishable from <span style="font-style: italic;">Erlicosaurus andrewsi</span> and <span style="font-style: italic;">Segnosaurus ghalbiensis</span>
bones in the collections of the Paleontological Institute (at the
Central State Museum) in Ulaan Baatar. There are a few elements from
the Iren Dabasu that may also be referable to the more poorly known
segnosaur <span style="font-style: italic;">Enigmosaurus</span>." They listed both <span style="font-style: italic;">Segnosaurus</span> sp. and <span style="font-style: italic;">Erlikosaurus</span>
sp. as present, but given the poor state of knowledge of therizinosaur
diversity in the early 90s, they may actually belong to the
contemporaneous <i>Erliansaurus,
Neimongosaurus</i>, and/or the
taxon to which the forelimb AMNH 6368 belongs instead. Precise
localities have not been published, but the CCDP excavated several
(Currie and Eberth, 1993: Table 3), all around Iren Nor and far from
Sanhangobi where <span style="font-style: italic;">Erliansaurus</span> and <span style="font-style: italic;">Neimongosaurus</span> were found.<br />
<b>References</b>- Dong, 1992. Dinosaurian Faunas of China: China Ocean Press. 188 pp.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People’s Republic
of China. Cretaceous Research. 14, 127-144.<br />
Dong, 1993. The field activities of the Sino-Canadian Dinosaur Project
in China, 1987-1990. Canadian Journal of Earth Sciences. 30(10),
1997-2001.</p><p><i><b>Kuszholia sp.</b></i> (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)<br />
<b>Middle-Late Campanian, Late Cretaceous<br />
Iren Dabasu Formation, Inner Mongolia, China</b><br />
<b>Material</b>- (IVPP V20377) anterior dentaries (symph 8.0 mm)<br />
<b>Comments</b>-
This was discovered in 2012 at "a rare microvertebrate locality within
the Iren Dabasu Formation, about 16 km northeast of Erenhot City",
which would put it in localities Q-T of Xing et al. (2012). Yao et al.
(2015) referred IVPP V20377 to <i>Caenagnathasia</i>
sp., and it does possess all characters here listed as diagnostic for
that taxon. Of their characters listed as varying between <span style="font-style: italic;">Caenagnathasia</span> specimens, most also vary between <span style="font-style: italic;">Chirostenotes</span>
specimens (posterior surface of symphysis with tubercle; chin-like
eminence between anterior and ventral surfaces; pneumatic foramen in
front of mandibular fenestra on lateral surface of dentary; depression
on posteroventral margin of symphysis [ontogenetic?]), while paired
second anterior occlusal grooves flanking first anterior occlusal
groove is polymorphic in the holotype, and lateral projections on
lingual ridges may be absent in ZIN PH 2354/16 due to preservation.
This leaves presence of a median symphyseal groove on the posterodorsal
depression in the holotype and IVPP V20377 but not ZIN PH 2354/16
(ontogenetic?) and on the posteroventral symphysis of IVPP V20377 but
not the holotype or ZIN PH 2354/16 (taxonomic?). The age difference
suggests the specimen is not conspecific with the Bissekty species.<br />
<b>References</b>-
Xing, He, Li and Xi, 2012. A review on the study of the stratigraphy,
sedimentology, and paleontology of the Iren Dabasu Formation, Inner
Mongolia. In Dong (ed.). Proceedings of the Thirteenth Annual Meeting
of the Chinese Society of Vertebrate Paleontology. China Ocean Press.
1-44.<br />
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. <i>Caenagnathasia</i>
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.<br />
Wang, Zhang and Yang, 2018. Reevaluation of the dentary structures of
caenagnathid oviraptorosaurs (Dinosauria, Theropoda). Scientific
Reports. 8:391. <br /></p><p><i><b>Avimimus sp. nov.</b></i> (Makovicky, 1995)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China</b><br />
<b>Material</b>- (AMNH 6570 in part; paratype of <span style="font-style: italic;">Ornithomimus asiaticus</span>) fibula (Chiappe, Norell and Clark, 2002)<br />
(AMNH 6576 in part; paratype of <span style="font-style: italic;">Ornithomimus asiaticus</span>) proximal caudal vertebra (Makovicky, 1995)<br />
?(AMNH 6754) distal metatarsal III (Ryan, Currie and Russell, 2001)<br />?..?(AMNH
6755; 'AMNH 6555' of Funston, Currie, Ryan and Dong, 2019) incomplete
metatarsus (mtII 100, mtIV 99 mm) (Ryan, Currie and Russell, 2001)<br />
(AMNH 25569) caudal vertebra, ten vertebrae, three phalanges, five unguals including pedal ungual (~18 mm) (AMNH online)<br />
(AMNH coll.) dorsal vertebrae (Makovicky, 1995)<br />
?(IVPP V16313.a) manual ungual ?I (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16314; = TMP 1992.302.0102) proximal tarsometatarsus (Ryan, Currie and Russell, 2001)<br />
?...(IVPP V16341) tarsometatarsus (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16315) metatarsal II (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16316.a) pedal ungual III (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16316.b) pedal ungual III (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16316.c; = TMP 1992.302.0119A) pedal ungual II/IV (Ryan, Currie and Russell, 2001)<br />
(IVPP V16316.d; = TMP 1992.302.0119B) pedal ungual II/IV (Ryan, Currie and Russell, 2001)<br />
(IVPP V16317.a) incomplete proximal caudal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16317.b) mid caudal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16318; ?= TMP 1992.302.0344) (juvenile) posterior cervical vertebra (Ryan, Currie and Russell, 2001)<br />
(IVPP V16318.a) incomplete anterior dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16318.b; ?= IVPP 160788-124) incomplete posterior dorsal vertebra (Makovicky, 1995; Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16319) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16320) (<1 year old juvenile) distal tibiotarsus (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16321) proximal metatarsal II (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16322.a; = TMP 1992.302.0150) proximal tibia (Ryan, Currie and Russell, 2001)<br />
(IVPP V16322.b) proximal tibia (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16322.c) proximal tibia (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16323.a) third dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16324) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16325) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16326) metatarsal II (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16327; = TMP 1992.302.0116) partial scapulocoracoid (Ryan, Currie and Russell, 2001)<br />
(IVPP V16328) (juvenile) mid sacral centrum (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16329.a; = IVPP 180788-123) last or penultimate cervical vertebra (Makovicky, 1995)<br />
(IVPP V16329.b) second dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16330; = IVPP 160788-122) posterior sacral fragment (Makovicky, 1995)<br />
(IVPP V16331) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16332.a; = TMP 1992.302.0140) partial second dorsal vertebra (Ryan, Currie and Russell, 2001)<br />
(IVPP V16332.b) partial first dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16333) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16334.a; = TMP 1992.302.0149) proximal femur (Ryan, Currie and Russell, 2001)<br />
(IVPP V16335.a) (juvenile) distal tarsal IV fused to proximal metatarsal IV (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16335.b) distal metatarsal II/III<br />
....(IVPP V16335.c) distal metatarsal II/III<br />
....(IVPP V16336) distal metatarsal IV <br />
(IVPP V16337) (>2 year old adult) distal tarsometatarsus (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16338; = TMP 1992.302.0110) distal femur (Ryan, Currie and Russell, 2001)<br />
(IVPP V16339) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16340; = TMP 1992.302.0117) proximal humerus (Ryan, Currie and Russell, 2001)<br />?(IVPP V16342; = TMP 1992.302.0104) partial frontal (Ryan, Currie and Russell, 2001)<br />
(IVPP V16343) distal metacarpal I (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16344) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V16345) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP V163...) four dorsal vertebrae, distal metatarsal, pedal ungual
III, eight pedal unguals II/IV (Makovicky, 1995; Funston, Currie, Ryan
and Dong, 2019)<br />
(PIN coll.) material (Currie and Eberth, 1993)<br />
<span style="font-weight: bold;">Diagnosis</span>- (after Funston et al., 2019) three cervicodorsal vertebrae (unknown in <span style="font-style: italic;">A. nemegtensis</span>); deeper femoral intercondylar groove.<br />
<b>Comments</b>-
Makovicky (1995) stated "Avimimid vertebrae were collected by the
American Museum Central Asian expedition in 1922 at Iren Dabasu.
However, they were not recognised as such, and were catalogued with <span style="font-style: italic;">Ornithomimus asiaticus</span> (now <span style="font-style: italic;">Archaeornithomimus asiaticus</span>) material", but the <span style="font-style: italic;">Archaeornithomimus</span>
type material was discovered in 1923 including a proximal caudal
referenced by the author. While he says "one of the caudals collected
by the American Museum's Central Asiatic Expeditions (AMNH 6576), [has]
a small pointed tubercle separates the ventromedial corners of the
chevron facets" this specimen number includes almost a hundred elements
from the Johnson Quarry AMNH locality 141 generally referred to <span style="font-style: italic;">Archaeornithomimus</span>.
A fibula (AMNH 6570) was figured as "alvarezsaurid fibula from Iren
Dabasu (Inner Mongolia, China)" by Chiappe et al. (2002), but Longrich
and Currie (2009) stated "it more closely resembles the fibula of
Avimimidae, which are common at this locality (N.R.L., pers. obs.)."
Again this number includes over two hundred paratype <span style="font-style: italic;">Archaeornithomimus</span>
elements, this time from the Kaisen Quarry AMNH locality 140.
Makovicky also stated "A large number of dorsal vertebrae are present
in the American Museum ... collections from Iren Dabasu", which are
likely to also be catalogued under AMNH 6570 and/or 6576. Longrich's
claim is supported here however as the fibula has an anteriorly
projected iliofibularis tuber as in <span style="font-style: italic;">Avimimus</span>
but unlike the laterally directed tuber of e.g. IGM 100/99 and is less
reduced distally in anteroposterior width than the latter. A distal
caudal
(AMNH coll.) described by Makovicky "has the morphology of a typical
coelurosaurian
distal caudal, but is otherwise undiagnostic. The possibility that it
may originate from an avimimid is suggested by its small size. It
should be noted, however, that it could just as conceivably be from the
tail of a juvenile <span style="font-style: italic;">Archaeornithomimus</span> from the same bonebed." Indeed, the <span style="font-style: italic;">A. nemegtensis</span> bonebed show <span style="font-style: italic;">Avimimus</span> has short distal caudals like other caenagnathoids and unlike ornithomimosaurs, so this specimen is here referred to <span style="font-style: italic;">Archaeornithomimus</span>.
The AMNH online catalogue lists AMNH 25569 as "10 vertebrae 1 caudal
vertebra 3 phalanges & 5 claws" of Saurischia from the Johnson
Quarry with a pedal ungual photographed as Avimimidae. A metatarsus
(AMNH 6755) and third metatarsal (AMNH 6754) were listed on the
museum's online catalogue as <i>Elmisaurus sp</i>., but also referred to <i>Avimimus</i> by Ryan et al. (2001) and Funston et al. (for AMNH 6755 at least)<span style="font-style: italic;"></span>.
Ryan et al.'s poster indicated both were found in 1923 and were
possibly associated. AMNH 6755 does seem smaller than other fused <span style="font-style: italic;">Avimimus</span>
and less slender, so further study is necessary. Note Funston et al.
(2019) describe and figure this as AMNH 6555, which is the number of
ornithischian material. Funston et al. state the main Iren Dabasu <span style="font-style: italic;">Avimimus</span>
bonebed "was originally discovered by a Sino-Soviet expedition in 1959,
which used bulldozers to excavate the site. It was revisited in 1987
and 1988 by the Sino-Canadian expedition and numerous fragmentary bones
representing all regions of the skeleton (Fig. 2) were recovered from
the spoil piles left by the Sino-Soviet bulldozers. Unfortunately, the
material collected by the Sino-Soviet expedition still awaits
preparation and it may never be available for study." This indicates
it must be locality K of Currie and Eberth (1993), which was CCDP
locality 1 and may correspond to AMNH locality 141. One element from
this excavation was described and figured by Kurzanov (1987) as "a left
avimimid femur from the Upper Cretaceous Iren-Nor locality in China
(specimen PIN, no. 2549-100)" (translated) but is here placed in
Oviraptoridae. While the PIN Sino-Soviet <span style="font-style: italic;">Avimimus</span>
material remains undescribed (although mentioned by Currie and Eberth),
the Sino-Canadian material was noted by Dong et al. (1989) who reported
"seven pedal elements of
<span style="font-style: italic;">Avimimus</span>" discovered in July 1988 and Dong (1992) refers to
"fused tarsometatarsi of <span style="font-style: italic;">Avimimus</span>" recovered on that expedition in July 1998.
Currie and Eberth (1993) stated "Direct comparison between isolated <span style="font-style: italic;">Avimimus</span> bones from the Iren Dabasu and the type specimen of <span style="font-style: italic;">Avimimus portentosus</span>
in Moscow failed to reveal any differences" and said the "material is
presently under review (Currie, Zhao and Kurzanov, in preparation)."
The Sino-Canadian <span style="font-style: italic;">Avimimus</span> material was eventually described by Makovicky, Ryan et al., then officially by Funston et al. as <span style="font-style: italic;">Avimimus</span>
sp.. Makovicky used IVPP field numbers (of which IVPP 180788-123 may
be a mistake for IVPP 160788-123 to better match the other two listed
numbers and only differ in the last digit), Ryan et al. used TMP
numbers in their poster, and the specimens were seemingly eventually
transported back to the IVPP for permanent storage. Note the "isolated
left frontal" mentioned by Sues et al. is not described in Funston et
al. (although the frontal is highlighted as preserved in their figure
2), but Funston (2019) describes it in his thesis chapter that was
developed into that paper. In it he says "whether it pertains to an
avimimid or another oviraptorosaur is uncertain. It can be
distinguished from other theropods by the large, incising nasal
contact, which is similar to the morphology of <span style="font-style: italic;">Elmisaurus rarus</span>",
so it's possible this belonged to the Iren Dabasu caenagnathid taxon
represented by dentaries IVPP V20377. IVPP 160788-124 is one of three
posterior dorsals without a ventral keel, so may be IVPP V16318.b which
was the only one of these figured by Funston et al.. Another tricky
specimen is TMP 1992.302.0344 as illustrated in Ryan et al.'s poster,
which is tentatively identified as posterior cervical IVPP V16318 as it
seems to have a ventrally placed parapophysis, an elongate centrum with
vertical articular surfaces, two or three central foramina and a dark
neurocentral boundary perhaps representing an open suture, but contra
to Funston et al.'s description is not notably smaller than other
cervicals. While Funston et al. only referred to the Iren Dabasu
material as Avimimidae gen. et sp. indet., they also stated "the
cervicodorsal vertebrae differ in number (three with hypapophyses) from
those of <span style="font-style: italic;">Avimimus portentosus</span> (MPC-D 100/129), although cervicodorsal number in <span style="font-style: italic;">Avimimus nemegtensis</span>
is unknown. The distal condyles of the femur (Fig. 6) are separated
much more deeply than is typical in avimimids, and metatarsals II and
IV (Fig. 8) are much more disparate in size", but ended up concluding
"the available material from the Iren Dabasu bonebed is too incomplete
to confidently erect a new taxon, but future
preparation of the Russian material (or collection of new material) may
result in its taxonomic distinction from other avimimids."<br /><b>References</b>- Kurzanov, 1987. Avimimidae and the problem of the origin
of birds. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa
ekspedicia. 31, 1-95.<br />
Dong, Currie and Russell, 1989. The 1988 field program of the Dinosaur Project. Vertebrata Palasiatica. 27(3), 233-236.<br />
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People's Republic
of China. Cretaceous Research. 14, 127-144.<br />
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp. <br />
Ryan, Currie and Russell, 2001. New material of <i>Avimimus portentosus</i>
(Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the
Erenhot region of Inner Mongolia. Journal of Vertebrate Paleontology.
21(3), 95A.<br />
Chiappe, Norell and Clark, 2002. The Cretaceous, short-armed Alvarezsauridae,
<i>Mononykus</i> and its kin. In Chiappe and Witmer (eds.). Mesozoic
Birds: Above the Heads of Dinosaurs. University of California Press. 87-120.<br />
Longrich and Currie, 2009 (online 2008). <i>Albertonykus borealis</i>, a new alvarezsaur (Dinosauria:
Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for
the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1),
239-252.<br />
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.<br />
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age
flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports.
9:18816. </p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjemw29kHrV5zIIWnAb1yDPTbsPDdF6CmSJhnlQj4XlgcqAATaPPXRhGZJdgvinkpvG6ydOB2ogvLOnooqekdz3dfXFAXtw5Ycg4oE7vWM9wEljp41XffLiJzIiz1O8ttho48aLSBZcqU2-_k_yWx3ljiiySRGPDUokzUa688ILSj1XSuDw_BaDYIqE/s2048/Archaeornithomimus%20AMNH%206570%20drawer~5.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1536" data-original-width="2048" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjemw29kHrV5zIIWnAb1yDPTbsPDdF6CmSJhnlQj4XlgcqAATaPPXRhGZJdgvinkpvG6ydOB2ogvLOnooqekdz3dfXFAXtw5Ycg4oE7vWM9wEljp41XffLiJzIiz1O8ttho48aLSBZcqU2-_k_yWx3ljiiySRGPDUokzUa688ILSj1XSuDw_BaDYIqE/s320/Archaeornithomimus%20AMNH%206570%20drawer~5.JPG" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Small portion of material catalogued under AMNH 6570 and assigned to <i>Archaeornithomimus</i>. Note the ungual in the upper left is not ornithomimid and here compared favorably to <i>Citipati</i>. My photo courtesy of the AMNH.<br /></td></tr></tbody></table><br /></p><p><span style="font-weight: bold;">cf. <span style="font-style: italic;">Citipati</span></span> (Kurzanov, 1987)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material-</b> ?(AMNH 6570 in part; paratype of <span style="font-style: italic;">Ornithomimus asiaticus</span>) partial manual ungual ?I (~39 mm) (pers. obs.)<br />
(PIN 2549-100) femur (~160 mm) (Kurzanov, 1987)<br />
<b>Comments-</b> Kurzanov (1987) briefly described and figured "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="ru" data-number-of-phrases="1" data-phrase-index="0"><span class="Q4iAWc">a
left avimimid femur from the Upper Cretaceous Iren-Nor locality in
China (specimen PIN, no. 2549-100)" (translated), which Currie and
Eberth (1993) indicate was found by the Sino-Soviet expedition, which means it was collected between </span></span></span>June 14 and July
17 at their localities K (= AMNH locality 141?), L or P. Kurzanov
referred it to Avimimidae based on the "accessory condyle" (which is
just the lateral condyle being separated from the ectocondylar tuber by
a fibular groove as in most theropods), the broad intercondylar flexor
groove and similarities of the trochanteric crest. Indeed, he stated "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="ru" data-number-of-phrases="1" data-phrase-index="0"><span class="Q4iAWc">the
only not very significant difference is expressed in the fusion of the
large and small trochanters, while in <span style="font-style: italic;">Avimimus</span> they are separated by a
narrow gap." On the other hand, Osmolska (1996) stated "There is a
great resemblance between the femur in<span style="font-style: italic;"> </span>[<span style="font-style: italic;">Bagaraatan</span>]<span style="font-style: italic;"> ostromi</span> and the femur PIN
2549-100" in that "Both femora have similarly shaped proximal and
distal ends, ... well pronounced articular heads and femoral necks, the
poorly delimited lesser trochanters, which are as high as the greater,
and in the presence of the protuberances on the lateral surface."
Additionally, "The distal ends of femora are also similarly shaped in
both compared forms" with "distinctive tibiofibular crests ('condylus
lateralis' in Kurzanov 1987)." While PIN 2549-100 is similar in shape
to both Iren Dabasu <span style="font-style: italic;">Avimimus</span> and <span style="font-style: italic;">Bagaraatan</span>, the latter both have
accessory trochanters (usually misidentified as a large, distally
placed anterior trochanter in <span style="font-style: italic;">Avimimus</span>) which are absent in PIN
2549-100, <span style="font-style: italic;">Avimimus</span> differs from PIN 2549-100 and <span style="font-style: italic;">Bagaraatan</span> in lacking
a distal ectocondylar notch defining the tuber, <span style="font-style: italic;">Bagaraatan</span> differs from
PIN 2549-100 and <span style="font-style: italic;">Avimimus</span> in having a narrow flexor groove, and PIN
2549-100 differs from at least <span style="font-style: italic;">Avimimus</span> in having a fourth trochanter
reduced to "</span></span></span><span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="ru" data-number-of-phrases="1" data-phrase-index="0"><span class="Q4iAWc">a
slight roughness, located almost under the head of the femur on its
medial side" (Kurzanov, 1987) (unknown in <span style="font-style: italic;">Bagaraatan</span>). Currie and
Eberth (1993) believed PIN 2549-100 "is probably from a troodontid" and
"provisionally referred to <span style="font-style: italic;">Saurornithoides</span>", but Averianov and Sues
(2012) concluded it "is probably troodontid but cannot be definitely
referred to <span style="font-style: italic;">Saurornithoides</span>" and "should be listed as Troodontidae
indet." However, scoring this in Hartman et al.'s maniraptoromorph analysis results in identical scorings to <span style="font-style: italic;">Citipati osmolskae</span>, with one more step needced to move it sister to <span style="font-style: italic;">Avimimus</span> and two more to move it to Troodontidae (as sister to <span style="font-style: italic;">Linhevenator</span>). Given the stratigraphic and geographic proximity, it is provisionally assigned to cf. <span style="font-style: italic;">Citipati</span> here pending description of the holotype's femur (IGM 100/979 and 1004 have crushed and poorly exposed femora).<br />
</span></span></span>The hypodigm of <span style="font-style: italic;">Archaeornithomimus asiaticus</span> includes two collections
of largely undescribed and unassociated specimens, AMNH 6570 from Third
Asiatic Field site 140 and AMNH 6576 from site 141, discovered between April 22 and May 25 1923. Based on personal
examination (July 2009), multiple elements in these collections do not
belong to <span style="font-style: italic;">Archaeornithomimus</span>, among which is an ungual in a box of
phalanges and calcanea under AMNH 6570. It is moderately curved with a
large, proximally placed flexor tubercle and resembles both
<span style="font-style: italic;">Sinornithoides</span>' pedal ungual I and <span style="font-style: italic;">Citipati</span>'s
manual ungual I in its
preserved portion. As it is twice the length of Iren Dabasu's
troodontid pedal unguals I but scales well to PIN 2549-100, the latter
identification is provisionally preferred here.<br />
<b>References</b>- Kurzanov, 1987. Avimimidae and the problem of the origin
of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa
ekspedicia. 31, 1-95.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and
palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia,
People's Republic of China. Cretaceous Research. 14, 127-144.<br />
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt
Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.<br />
Averianov and Sues, 2012. Correlation of Late Cretaceous continental vertebrate
assemblages in middle and central Asia. Journal of Stratigraphy. 36(2), 462-485.</p><p><b>undescribed Troodontidae</b> (Currie and Eberth,
1993)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>Erenhot, Iren Dabasu Formation, Inner Mongolia, China</b><br />
<b>Material</b>- (AMNH 6570 in part; paratype of <span style="font-style: italic;">Ornithomimus asiaticus</span>) (juvenile or subadult) axis, third cervical vertebra, fifth cervical vertebra (Makovicky,
1995)<br />
?(AMNH 6576 in part; paratype of <span style="font-style: italic;">Ornithomimus asiaticus</span>) pedal ungual I (~17 mm) (pers. obs.)<br />
(AMNH 21751) distal metatarsals III (Currie and Eberth,
1993)<br />
(AMNH 21772) metatarsal II (~183 mm) (Currie and Eberth, 1993)<br />
(AMNH 25570) three vertebrae (AMNH online)<br />
(AMNH 30261) proximal metatarsal (AMNH online)<br />
(AMNH 30262) proximal tibial fragment (AMNH online)<br />
(AMNH 30263) proximal tibial fragment (AMNH online)<br />
(AMNH 30264) tibial fragment, fibular fragment (AMNH online)<br />
(AMNH 30265) proximal tibia (AMNH online) 142<br />
(AMNH 30266) proximal fibula (AMNH online) 142<br />
(AMNH 30267) proximal tibia (AMNH online)<br />
(AMNH 30268) proximal fibula (AMNH online)<br />
(AMNH 30269) proximal fibula (AMNH online) 142<br />
(AMNH 30270) proximal fibula (AMNH online)<br />
(AMNH 30271) partial astragalus (AMNH online)<br />
(AMNH 30272) partial astragalus (AMNH online)<br />
(AMNH 30273) partial astragalus (AMNH online)<br />
(AMNH 30274) partial astragalus (AMNH online)<br />
(AMNH 30275) distal humerus (AMNH online)<br />
(AMNH 30276) distal humerus (AMNH online)<br />
(AMNH 30277) distal humerus (AMNH online)<br />
(AMNH 30278) proximal humerus (AMNH online)<br />
(AMNH 30279) proximal humerus (AMNH online)<br />
(AMNH 30280) proximal ulna (AMNH online)<br />
(AMNH 30281) distal radius (AMNH online)<br />
(AMNH 30282) distal radius (AMNH online)<br />
(AMNH 30283) proximal scapula (AMNH online)<br />
(AMNH 30284) proximal scapula (AMNH online)<br />
(AMNH 30285) scapular blade (AMNH online)<br />
(AMNH 30286) pedal ungual I (~19 mm) (AMNH online)<br />
(AMNH 30287) proximal manual ungual (AMNH online)<br />
(AMNH 30288) two posterior cervical or proximal caudal vertebrae (AMNH online)<br />
(AMNH 30289) distal metatarsal IV (AMNH online)<br />
(AMNH 30290) distal metatarsal IV (AMNH online)<br />
(AMNH 30291) distal metatarsal IV (AMNH online)<br />
(AMNH 30292) distal metatarsal III (AMNH online)<br />
(AMNH 30293) distal metatarsal III (AMNH online)<br />
(AMNH 30294) distal metatarsal II (AMNH online)<br />
(AMNH 30295) distal metatarsal II (AMNH online)<br />
(AMNH 30296) distal metatarsal II (AMNH online)<br />
(AMNH 30297) distal metatarsal II (AMNH online)<br />
(AMNH 30300) partial ilium (AMNH online) 142<br />
(AMNH 30301) proximal ?pubis (AMNH online) 142<br />
(AMNH 30302) ?ilial fragment (AMNH online)<br />
(AMNH 30303) partial synsacrum (AMNH online) 142<br />
(AMNH 30304) proximal ?pubis (AMNH online)<br />
?(AMNH 30305) last sacral vertebra (~25 mm) (AMNH online)<br />
(AMNH 30306) partial synsacrum (AMNH online)<br />
(AMNH 30307) synsacral fragment(AMNH online)<br />
(AMNH 30308) partial posterior cervical vertebra (AMNH online)<br />
(AMNH 30309) partial posterior cervical vertebra (AMNH online)<br />
(AMNH 30310) partial anterior dorsal vertebra (AMNH online)<br />
(AMNH 30311) partial anterior dorsal vertebra (AMNH online)<br />
(AMNH 30312) partial anterior dorsal centrum (AMNH online)<br />
(AMNH 30313) incomplete anterior dorsal centrum (AMNH online)<br />
(AMNH 30314) partial anterior dorsal centrum (AMNH online)<br />
(AMNH 30315) incomplete anterior dorsal centrum (AMNH online)<br />
(AMNH 30316) partial anterior dorsal vertebra (AMNH online)<br />
(AMNH 30317) incomplete anterior dorsal centrum (~24 mm) (AMNH online)<br />
(AMNH 30318) anterior dorsal centrum (~26 mm) (AMNH online)<br />
(AMNH 30320) anterior dorsal centrum (~26 mm) (AMNH online) 142<br />
(AMNH 30321) partial anterior dorsal vertebra (~26 mm) (AMNH online)<br />
?(AMNH 30322) anterior dorsal centrum (~32 mm) (AMNH online)<br />
(AMNH 30323) incomplete posterior dorsal centrum (~22 mm) (AMNH online)<br />
(AMNH 30324) incomplete dorsal centrum (AMNH online)<br />
(AMNH 30325) posterior dorsal centrum (~26 mm) (AMNH online)<br />
(AMNH 30326) posterior dorsal centrum (~27 mm) (AMNH online)<br />
(AMNH 30327) posterior dorsal centrum (~27 mm) (AMNH online)<br />
(AMNH 30328) incomplete posterior dorsal vertebra (~27 mm) (AMNH online)<br />
(AMNH 30329) incomplete posterior dorsal vertebra (~29 mm) (AMNH online)<br />
(AMNH 30330) proximal caudal centrum (~23 mm) (AMNH online)<br />
(AMNH 30336) ?central fragment (AMNH online)<br />
(AMNH 30337) incomplete distal caudal vertebra (~35 mm) (AMNH online)<br />
(AMNH 30338) mid caudal vertebra (~32 mm) (AMNH online)<br />
(AMNH 30339) incomplete mid caudal vertebra (~31 mm) (AMNH online)<br />
(AMNH 30340) incomplete mid caudal vertebra (AMNH online)<br />
(AMNH 30341) partial distal caudal vertebra (AMNH online)<br />
(AMNH 30342) mid caudal vertebra (~29 mm) (AMNH online)<br />
(AMNH 30343) mid caudal vertebra (~30 mm) (AMNH online)<br />
(AMNH 30344) distal caudal vertebra (~32 mm) (AMNH online)<br />
(AMNH 30345) incomplete mid caudal vertebra (~25 mm) (AMNH online)<br />
(AMNH 30346) partial distal caudal vertebra (AMNH online)<br />
(AMNH 30347) partial distal caudal vertebra (AMNH online)<br />
(AMNH 30348) distal caudal vertebra (~31 mm) (AMNH online)<br />
(AMNH 30349) fragmentary distal caudal vertebra (AMNH online)<br />
(AMNH 30350) distal caudal vertebra (~33 mm) (AMNH online)<br />
(AMNH 30351) incomplete distal caudal vertebra (~31 mm) (AMNH online)<br />
(AMNH 30352) partial distal caudal vertebra (AMNH online)<br />
(AMNH 30353) incomplete distal caudal vertebra (AMNH online)<br />
(AMNH 30354) distal caudal vertebra (~23 mm) (AMNH online)<br />
?(AMNH 30355) mid caudal vertebra (~32 mm) (AMNH online)<br />
(AMNH 30356) proximal caudal centrum (~28 mm) (AMNH online)<br />
(AMNH 30357) incomplete proximal caudal vertebra (~25 mm) (AMNH online)<br />
(AMNH 30358) proximal caudal vertebra (~24 mm) (AMNH online)<br />
(AMNH 30359) incomplete proximal caudal centrum (AMNH online)<br />
(IVPP 230790-16; = IVPP 230090-16 of Currie and Eberth, 1993) metatarsal III (Currie and Eberth, 1993)<br />
<b>Comments-</b>
Currie and Eberth (1993) stated "Troodontid bones are rare, but include
distinctive third metatarsals (AMNH 21751, 21772, IVPP 230090-16), in
which the distal articulation extends onto the posterior surfrace of
the bone in a broad tongue." However, Currie and Dong (2001) corrected
the identification of the second specimen, stating "AMNH 21772 is the
proximal end of a second metatarsal. It is identified as a troodontid
on the basis of its contact surface for the fourth metatarsal, its
size, and especially its lateromedial compression." The AMNH online
catalogue photo indicates most of the element is preserved and the
locality info is "8 mi. E. of station" indicating it was found in
localities 140-149 in 1923 or 1928. Currie and Dong describe AMNH
21751 as "two distal ends of third metatarsals [that] are about the
same size and represent left and right elements. Although they may
represent the same individual, the two fossils are different colours,
which suggests they may not have been found together." They indicate
these were "Collected in the 1920s by the third Central Asiatic
Expedition from exposures of the Iren Dabasu Formation (?Santonian)
near Erenhot." Currie and Dong list IVPP 230790-16 (presumably the
correct field number for Currie and Eberth's 'IVPP 230090-16') as a
metatarsal "Collected in 1990 from exposures of the Iren Dabasu
Formation (?Santonian) near Erenhot", which would make it found during
the second Sino-Canadian expedition. They state "the tongue-like
extensions of the third metatarsals from Iren Dabasu are flat like
those of <span style="font-style: italic;">Troodon</span> ... , <span style="font-style: italic;">Borogovia</span> ... , and <span style="font-style: italic;">Tochisaurus</span>" but unlike the grooved surface of <span style="font-style: italic;">Sinornithoides</span> or the distally restricted surface of <span style="font-style: italic;">Philovenator</span>. This has since been identified in Bissekty <span style="font-style: italic;">Urbacodon</span> sp. ZIN PH 2342/16, and it should be noted the extension of <span style="font-style: italic;">Tochisaurus</span> is much shorter, while <span style="font-style: italic;">Mei</span>, IGM 100/44, 100/140 and 100/1126 have a condition like <span style="font-style: italic;">Sinornithoides</span>. Thus as hypothesized by Dong and Currie, at least AMNH 21751 and IVPP 230790-16 are closer to <span style="font-style: italic;">Troodon</span> than <span style="font-style: italic;">Sinornithoides</span>. Currie and Eberth stated "These bones are <span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="ru" data-number-of-phrases="1" data-phrase-index="0"><span class="Q4iAWc">provisionally referred to <span style="font-style: italic;">Saurornithoides</span>" (at the time a concept including <span style="font-style: italic;">Zanabazar</span>)
without rationale, but Currie and Dong instead classified them as "an
unknown species of troodontid", stating they "cannot be identified
further without additional material."</span></span></span><span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="ru" data-number-of-phrases="1" data-phrase-index="0"><span class="Q4iAWc"></span></span></span><br />
Makovicky (1995) stated "A probable troodontid axis (AMNH 6570),
articulated with a third cervical vertebra, is present in the
collections of the American Museum of Natural History. This
identification is based on the morphology of the associated third
cervical and a probable fifth cervical, possibly from the same
individual, which strongly resembles those of <span style="font-style: italic;">Troodon</span>.
The axis is from an immature individual as seen from absence of both
the odontoid and axial intercentrum." This specimen number includes
over two hundred paratype <span style="font-style: italic;">Archaeornithomimus</span>
elements from the Kaisen Quarry AMNH locality 140, and the cervicals
described were not recognized in the material catalogued under it in
July 2009 (pers. obs.). However, a small ungual was noticed in AMNH
6576 (which includes almost a hundred paratype <span style="font-style: italic;">Archaeornithomimus</span>
elements from the Johnson Quarry AMNH locality 141) that most closely
resembles a troodontid pedal ungual I in the slight curvature,
proximally placed flexor tubercle and posterodorsal extent being less
than its posteroventral extent.<br />
The AMNH online catalogue lists AMNH 25570 as "<span style="font-style: italic;">Troodon ?</span>",
consisting of "3 vertebrae." A large number of elements (AMNH
30261-30297, 30300-30318, 30320-30330, 30336-30359) are labeled are
labeled "Troodontid" on the AMNH online catalogue, each from the same
location ("8 mi. E. of station") and from AMNH Quarry 142 specifically
when visible in the photo (AMNH 30265, 30266, 32069, 30300, 30301,
30303, 30320). Given the similar preservation and number of elements
preserved, it is possible these represent two individuals, and that
several other specimens only identified to the level of Saurischia in
the online catalogue (AMNH 30245, 30247-30260, 30298-30299, 30360) that
are also from "8 mi. E. of station" may belong to them as well. Note
AMNH 30267 is incorrectly identified as a proximal fibula, while 30288
is called "Proximal end of metatarsal IV" but seems to be two vertebrae
instead, AMNH 30301 is called an "Ilium fragment." but may be a
proximal pubis (posterior edge downward in photo), AMNH 30302 is
labeled as "Acetabulum fragment." and indeed may be the ischial
peduncle and postacetabular base of a left ilium, AMNH 30304 is labeled
"Prox. end of ischium" but more closely resembles a proximal troodontid
pubis in the diverging peduncles and shallowly concave acetabular edge,
AMNH 30305 is a last sacral vertebra with a convex posterior central
face and 30322 is an anterior dorsal with convex anterior central face
so both may be alvarezsaurid instead. Scoring the material as
photographed in the online catalogue (with AMNH 30305 and 30322
excluded, and 30301 and 30304 interpreted as pubes) into Hartman et
al.'s maniraptoromorph matrix does result in it being troodontid, but
note examination of the specimens themselves would provide far more
data for each element and that it's currently only an assumption that
they belong to the same taxon.<br />
<b>References</b>- Currie and Eberth, 1993. Palaeontology, sedimentology and
palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia,
People's Republic of China. Cretaceous Research. 14, 127-144.<br />
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp.<br />
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38(12), 1753-1766. <br />
Averianov and Sues, 2012. Correlation of Late Cretaceous continental vertebrate
assemblages in middle and central Asia. Journal of Stratigraphy. 36(2), 462-485.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh8xu1NyfUvmh2o6LTf948t-1OKma7l_Emd9VsnYPJrl4riPjuUCgzPTyRofl3I63w3UqVOfper73aHMUmBiZmqICJCscrkihxsxPIgVBluFt_YS2PO8nZ1dz-rDQgXgRpG4A0-JQZFjfpUs6eNRapftP7fpZRBqlBimNFtaTVC5JgNqMTm4jPTs6kh/s2048/Archaeornithomimus%20AMNH%206576~1.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1536" data-original-width="2048" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh8xu1NyfUvmh2o6LTf948t-1OKma7l_Emd9VsnYPJrl4riPjuUCgzPTyRofl3I63w3UqVOfper73aHMUmBiZmqICJCscrkihxsxPIgVBluFt_YS2PO8nZ1dz-rDQgXgRpG4A0-JQZFjfpUs6eNRapftP7fpZRBqlBimNFtaTVC5JgNqMTm4jPTs6kh/s320/Archaeornithomimus%20AMNH%206576~1.JPG" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Sample of unguals catalogued under AMNH 6576 as <i>Archaeornithomimus</i>. The bottom left one is not ornithomimid and is compared to pedal ugual I of <i>Sinornithoides </i>here, so suggested to be troodontid. Note too the juvenile ?<i>Bactrosaurus </i>pedal ungual in the box on the right. Scale = 100 mm. My photo courtesy of the AMNH.<br /></td></tr></tbody></table><br /><p><b>undescribed Dromaeosauridae</b> (Gilmore, 1933)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material</b>- (AMNH 6572) pedal phalanx II-1 (Ostrom, 1969)<br />
(AMNH 21781) pedal ungual II (AMNH online)<br />
?(IVPP 270790-4) tooth (~21x~9x? mm) (Currie and Zhao, 1993) <br />
?(IVPP V16334.b) proximal femur (Funston, Currie, Ryan and Dong, 2019)<br />
(IVPP coll.) teeth, elements (Dong, Currie and Russell, 1989)<br />
(IVPP coll.) teeth and/or elements (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)<br />
<b>Comments</b>-
Gilmore (1933) noted the presence of "a few foot bones and other
fragmentary skeletal parts" of small theropods from the Iren Dabasu
Formation that he assigned to "Dromaeosaurinae Genus and species
indet.", but admitted that this assignment as opposed to Coeluridae or
Compsognathidae is based purely on their Cretaceous age. Thus while it
turned out at least two of these elements are apparently dromaeosaurid
(see below), Gilmore's rationale is equivalent to Maniraptora indet.
today and the fossils referenced plausibly included material now
recognized as avimimid, troodontid and ?alvarezsaurid. Ostrom (1969)
noted in a discussion of deinonychosaur pedal examples that "E. H.
Colbert has also discovered an isolated phalanx (AMNH 6572) in the
American Museum collections from the Iren Dabasu Formation of Mongolia
which compares almost exactly with the proximal phalanx of digit II of <span style="font-style: italic;">Deinonychus</span>, but is perhaps 20 percent larger", which would make it somewhere around 46-52 mm long. He shows it questionably derived from <span style="font-style: italic;">Velociraptor</span>
in his phylogram without explanation, which would not make sense in the
most recent interpretation of Iren Dabasu's age being contemporaneous
or slightly older than the Djadochta. Paul (1988) states "at the AMNH
is a hyper-extendable toe bone from the Late Cretaceous of Mongolia
that looks like a <span style="font-style: italic;">Velociraptor</span> somewhat bigger than <span style="font-style: italic;">V. antirrhopus</span> [= <span style="font-style: italic;">Deinonychus</span>]",
but it is uncertain whether he saw it independant of Ostrom's text
(Paul, pers. comm. 6-2022). The AMNH online catalogue lists AMNH 21781
as an Iren Dabasu member of Dromaeosauridae represented by "Ungual of
pes (digit II)" and being found by Kaisen. Both AMNH 6572 and 21781
would have been found in Erenhot during the April 22 to May 25 1923
Central Asiatic Expedition.<br />Dong et al. (1989) state <span style="font-style: italic;">Velociraptor</span> material was discovered in the July 1988 Sino-Canadian expedition to Erenhot, and Dong (1992) specifies "teeth of <span style="font-style: italic;">Velociraptor</span>". Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in
the Iren Dabasu" and that "Most of these can be attributed to <i>Velociraptor</i>,
although some of the teeth suggest that there was a second, larger
species of an indeterminate dromaeosaurine dromaeosaurid." Yet no
rationale was presented, and the only two specified Iren Dabasu
dromaeosaurid elements in the literature are clearly not <i>Velociraptor</i> (AMNH 6572 is twice
the size, while IVPP 270790-4 is different from most dromaeosaurid
teeth as noted above). AMNH 6572 may belong to the supposed
dromaeosaurine though, based on size. Currie and Zhao (1993) figure
"Dromaeosaurid tooth (IVPP 270790-4) from
the Iren Dabasu Formation near Erenhot, People's Republic of China,
showing replacement pit on medial side of root", but given the slight
constriction basal to the
crown, convex distal edge and seeming lack of serrations, this may be
misidentified. Based on its similar field number to troodontid
metatarsal
IVPP 230790-16 it was probably also found in the Sino-Canadian
expedition of 1990.<br />
Funston et al. (2019) notes a femoral head that
supposedly differs from <span style="font-style: italic;">Avimimus</span>
in having fused anterior and greater trochanters, "which suggests that
it may be oviraptorid or, more likely, dromaeosaur." This is from the <span style="font-style: italic;">Avimimus</span>
bonebed (locality K of Currie and Eberth [1993], which was CCDP
locality 1 and may correspond to AMNH locality 141) and recovered in
July 1988. Interestingly, femur PIN 2549-100 is from the same locality
and shares a trochanteric crest, but is here identified as
oviraptorid. It's possible they are from the same taxon (or even
individual).<br />
Yao et al. (2015) note "small unarticulated bones and teeth, including
fossils of ... dromaeosaurids" from "a rare microvertebrate locality
within the Iren Dabasu Formation, about 16 km northeast of Erenhot
City."<br />
<b>References</b>- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.<br />
Ostrom, 1969. Osteology of <i>Deinonychus antirrhopus</i>, an unusual theropod
from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin.
30, 1-165.<br />
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
<br />
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.<br />
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic
of China. Cretaceous Research. 14, 127-144.<br />
Currie and Zhao, 1993 (published 1994). A new troodontid (Dinosauria, Theropoda) braincase from
the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth
Sciences. 30(10-11), 2234-2247.<br />
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. <i>Caenagnathasia</i>
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.<br />
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age
flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports.
9:18816.</p><p><b>undescribed possible dromaeosaurine</b> (Currie and Eberth, 1993)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material</b>- teeth<br />
<b>Comments</b>- Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in
the Iren Dabasu" and that "Most of these can be attributed to <i>Velociraptor</i>,
although some of the teeth suggest that there was a second, larger
species of an indeterminate dromaeosaurine dromaeosaurid." A pedal
phalanx II-1 (AMNH 6572) mentioned by Ostrom (1969) as being 20% larger
than <i>Deinonychus</i> may belong
to the same taxon, based on size.<br />
<b>References</b>- Ostrom, 1969. Osteology of <i>Deinonychus antirrhopus</i>,
an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of
Natural History Bulletin. 30, 1-165.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic
of China. Cretaceous Research. 14, 127-144.</p><p><b>undescribed Averostra</b> (Chow and Rozhdestvensky, 1960)<br />
<b>Middle-Late Campanian, Late Cretaceous</b><br />
<b>
Iren Dabasu Formation, Inner Mongolia, China<br />
Material</b>- (AMNH 6376) phalanx II-1 (AMNH online)<br />
(AMNH 6556) metatarsal II (AMNH online)<br />
(AMNH 6744) four caudal vertebrae, 8 distal pedal elements (AMNH online)<br />
(AMNH 6756) metatarsal (AMNH online)<br />
(AMNH 6757) limb fragments, metapodials, phalanx, fragments (AMNH online)<br />
(AMNH 21552) femur<br />
(AMNH 21565) elements<br />
(AMNH 21588) <br />
(AMNH 21774) fibula <br />
(AMNH 21775) pedal phalanx ?II-1<br />
(AMNH 21776) four proximal pedal phalanges <br />
(AMNH 21780) four unguals <br />
(AMNH 21782) manual ungual <br />
(AMNH 21784) four caudal vertebrae <br />
(AMNH 30245) two metatarsal II or IV shafts (AMNH online)<br />
(AMNH 30247) posterior dorsal rib fragment (AMNH online)<br />
(AMNH 30248) proximal anterior rib (AMNH online)<br />
(AMNH 30249) partial coracoid (AMNH online)<br />
(AMNH 30250) distal femur (AMNH online)<br />
(AMNH 30251) proximal femur (AMNH online)<br />
(AMNH 30252) distal femur (AMNH online)<br />
(AMNH 30253) proximal femur (AMNH online)<br />
(AMNH 30254) distal femur (AMNH online)<br />
(AMNH 30255) astragalus (AMNH online)<br />
(AMNH 30256) proximal tibia (AMNH online)<br />
(AMNH 30257) proximal femur (AMNH online)<br />
(AMNH 30258) distal tibia (AMNH online)<br />
(AMNH 30259) proximal metatarsal (AMNH online)<br />
(AMNH 30260) proximal metatarsal (AMNH online)<br />
(AMNH 30298) acetabular fragment (AMNH online)<br />
(AMNH 30299) proximal ischium (AMNH online)<br />
(AMNH 30360) metatarsal III shaft (AMNH online)<br />
(AMNH 80277) distal humerus (AMNH online)<br />
(IVPP or PIN coll.) (small) three partial skeletons (Chow and Rozhdestvensky, 1960)<br />
(IVPP and PIN coll.) <400 specimens (Currie and Eberth, 1993)<br />
<span style="font-weight: bold;">Comments</span>- The AMNH specimens
listed here are from the online catalogue, which generally lacks
identification for ranks between order and family so that Theropoda
indet. material is listed as Saurischia. Yet none of the specimens are
likely to be sauropods given Gilmore (1933) never mentioned finding any
and to this day only a few elements have been reported (4 in the
Erenhot Dinosaur Museum coll., 7 from the Sino-Soviet expedition-
Currie and Eberth, 1993; <span style="font-style: italic;">Sonidosaurus</span>).
Most would have been found during the April 22 to May 25 1923 Central
Asiatic Expedition, AMNH 6556 on April 30. The online catalog also
specifies AMNH 6756 was discovered in AMNH site 141. Many of these
specimens (AMNH 6556, 30245, 30247-30260, 30298-30299, 30360, 80277)
are listed as being from "8 mi. E. of station" which would place them
among Third Asiatic Expedition field sites 140-149, with AMNH 6757
listed as 9 miles east, so perhaps site 149. One exception is AMNH
6744, stated as being found at Elephant Camp (12 miles NW of the
station) by de Chardin, who was only on the 1930 expedition. Based on
the elements preserved, specimen numbers and locality of "8 mi. E. of
station", sevcral specimens (AMNH 30245, 30247-30260, 30298-30299,
30360) may belong to the two(+?) troodontid individuals noted in the
AMNH online catalog represented by specimen numbers AMNH 30261-30297,
30300-30318, 30320-30330 and 30336-30359.<br />
Chow and Rozhdestvensky (1960) noted "three partially complete
skeletons of some small carnosaurian dinosaurs" discovered in the
June-July 1959 Sino-Soviet expedition, perhaps indicating
tyrannosauroids or dromaeosaurids. Currie and Eberth (1993) stated "A
rough tally of Sino-Soviet field identifications shows that ...
'theropods' (including large theropods, small theropods and segnosaurs,
but not ornithomimids) were more common (400 specimens)."<br />
<span style="font-weight: bold;">References</span>- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.<br />
Chow and Rozhdestvensky, 1960. Exploration in Inner Mongolia - A
preliminary account of the 1959 field work of the Sino-Soviet
Plaeontological Expedition. Vertebrata PalAsiatica. 4(1), 1-10.<br />
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.<br />
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.<br />
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People’s Republic
of China. Cretaceous Research. 14, 127-144.<br />
Dong, 1993. The field activities of the Sino-Canadian Dinosaur Project
in China, 1987-1990. Canadian Journal of Earth Sciences. 30(10),
1997-2001. <br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com27tag:blogger.com,1999:blog-3248412803814730250.post-86374089699236558532022-05-24T04:54:00.001-07:002022-05-24T04:54:52.708-07:00Prehistoric Planet review<p>Back in 2011 I reviewed the big dinosaur documentaries of the time, Dinosaur Revolution and Planet Dinosaur. While both had the same basic concept to educate and entertain viewers about prehistoric life, they diverged wildly in style. <a href="https://theropoddatabase.blogspot.com/2011/09/dinosaur-revolution-review.html">Dinosaur Revolution</a> was "largely accurate-looking dinosaurs acting like humans in zany situations and learning valuable life lessons", while <a href="https://theropoddatabase.blogspot.com/2011/09/planet-dinosaur-review.html">Planet Dinosaur</a> was "generally inaccurate restorations behaving fairly realistically, packed full of references to specific discoveries in the literature, telling us what we know and why." Today I watched the first episode of the newest dinosaur documentary attempt, Prehistoric Planet, and figured I would continue the review tradition over a decade later.<br /><br />It's extremely good, a near perfect execution of the concept.<br /><br />I don't know if I've just gotten soft in my old age, but this was exactly what I wanted from a Walking With Dinosaurs type of show. The graphics are amazing, extremely close to the point where if I didn't know these animals were extinct I'd believe this was live footage. In fact, I'd believe the ammonites and pycnodonts were alive. My literal biggest complaint of the entire episode is that the <i>Tyrannosaurus</i>' skin didn't look like it was wet after emerging from the water, which was just several seconds of animation out of 42 minutes. While everything but the <i>Tyrannosaurus </i>was outside my area of expertise, I didn't see anything inaccurate or unrealistic as far as I could tell. <br /><br />As for the style of the show, all the animals acted like animals, which again is exactly what I want. On the one hand, behavior is ascribed to them that we have no direct evidence of in the fossil record, especially since most featured taxa (besides <i>Tyrannosaurus </i>and <i>Mosasaurus</i>) are poorly known. The Niobrara-esque pterosaur scenes are actually Couche III from the Maastrichtian of Morocco for instance, so use <i>Alcione</i>, <i>Tethydraco</i>, <i>Barbaridactylus</i>, etc.. But this works because the taxa themselves are never emphasized, and each behavior is plausible and only featured briefly until we move on to something else. '<i>Tuarangisaurus</i>' is treated just as the word 'rhinoceros' would be in a nature documentary without telling us what material it's known from or what country it was found in or anything geological at all. And we see them give live birth, display with upright necks out of water, swallow gastroliths, etc., but none of it is presented as essential to what <i>Tuarangisaurus </i>is or given a moral meaning to learn. It's just animal behaviors being done by animals, which is just what you get in nature documentaries that cover a wide array of species.<br /></p><p>Which combined with the inimitable David Attenborough (96 years old!) narrating makes this the best nature documentary-style dinosaur show. I don't want to see talking heads being edited to dumb down what I already know- if I want to learn about <i>Phosphatodraco</i>, I'll pull out Pereda-Suberbiola et al. 2003. But if I want the Maastrichtian to be filmed as if it still existed, Prehistoric Planet provides what the literature cannot.<br /><br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com3tag:blogger.com,1999:blog-3248412803814730250.post-17351441336335373702022-03-12T01:49:00.000-08:002022-03-12T01:49:50.264-08:00Theropod Database Update March 2022 with SVP 2021 thoughts<p>And after a few months we're back. This update incorporates taxa named since last time and also incorporates many of the theropod abstracts from SVP 2021.<br /><br />Two new ceratosaurs were added, <i><a href="https://www.theropoddatabase.com/Ceratosauria.htm#Guemesiaochoai">Guemesia</a> </i>and <a href="https://www.theropoddatabase.com/Ceratosauria.htm#Berthasauraleopoldinae"><i>Berthasaura</i></a>. The latter is interesting in just how similar to tetanurines, and coelurosaurs and ornithomimosaurs/oviraptorosaurs in particular, the supposed noasaurid is. Besides the obvious dentary similarities to e.g. <i>Microvenator </i>and highly pneumatic lacrimal, the typical ceratosaurian cervical postzygodiapophyseal lamina is reduced, the brevis fossa is reduced, and like <i>Elaphrosaurus </i>the femoral medial epicondyle is reduced and the fibular crest is separate from the proximal tibial condyles. The authors recover it as the most basal noasaurid, but this seems unlikely given the Aptian-Albian age.</p><p>Among basal tetanurines, <i><a href="https://www.theropoddatabase.com/Megalosauroidea.htm#Iberospinusnatarioi">Iberospinus</a> </i>and <a href="https://www.theropoddatabase.com/Coelurosauria.htm#Maipmacrothorax">"Maip"</a> were added, each formerly in the Database as a questionable referred specimen of <i>Suchosaurus? girardi</i> and Megaraptoridae indet.. <i><a href="https://www.theropoddatabase.com/Tyrannosauroidea.html#Dynamoterrordynastes">Dynamoterror</a> </i>was updated based on Yun's paper and the SVP abstract on the new specimen. I've incorporated Paul et al.'s new <i>Tyrannosaurus </i>species as synonyms, but haven't gone into any detail examining the issue yet.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/a/AVvXsEizKoks4gnMGhCzm2rZBlqZsEnRcZLYfuXp3FCmG6_R3QljyMvaHWVgCzB9ySY91Ofo5l3RuGo3m0LOwyLJZf5at2WEU9gZotdfIK3iOrwqEjeTG2rMDEkeTT3l7diWDAQQYODNxz4-y1pRfEBiZmC6KclHidRZD0SP9i-n1SoIk9cgQRHroSqcjhkT=s3072" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="2304" data-original-width="3072" height="240" src="https://blogger.googleusercontent.com/img/a/AVvXsEizKoks4gnMGhCzm2rZBlqZsEnRcZLYfuXp3FCmG6_R3QljyMvaHWVgCzB9ySY91Ofo5l3RuGo3m0LOwyLJZf5at2WEU9gZotdfIK3iOrwqEjeTG2rMDEkeTT3l7diWDAQQYODNxz4-y1pRfEBiZmC6KclHidRZD0SP9i-n1SoIk9cgQRHroSqcjhkT=s320" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Left ilium of <i>Ornitholestes hermanni</i> holotype AMNH 619 showing plesiomorphic characters not present in pennaraptorans- highly convex supracetabular crest and deep brevis fossa (courtesy of the AMNH).<br /></td></tr></tbody></table><i><br />Ornitholestes </i>was famously recovered as an oviraptorosaur by Chapelle et al. (2021), but while their CT scanning is amazing and I can't wait for the resulting paper, I'm skeptical of that phylogenetic placement. For one, some of their proposed synapomorphies are slightly closer to the oviraptorosaurian condition than neutral, but defining the state this loosely would result in other taxa also exhibiting the condition. The supposedly pendant paroccipital processes for instance are slightly downturned, but not nearly as much as oviraptorosaurs' and e.g. <i>Allosaurus </i>has a more extreme condition. They don't count as either short/deep or downturned using standard TWiG scoring or the Lori matrix. The other issue is that the postcranium is so plesiomorphic that basically all pennaraptoran characters become convergences between oviraptorosaurs and paravians, which might be true, but does keep an oviraptorosaurian <i>Ornitholestes </i>at 18 steps longer than the shortest trees in the Lori matrix even with the new data from CT scans. However, that data did lead to <i>Ornitholestes </i>moving crownward to be sister to Maniraptoriformes instead of the most basal maniraptoromorph.</p><p><a href="https://www.theropoddatabase.com/Coelurosauria.htm#Fukuivenatorparadoxus">"Fukuivenator"</a> just got a huge redescription (Hattori et al., 2021), with excellent CT scans in every view. They corrected most of what the original description got wrong (which was even more than I thought), and surprise(!) the weirdly large premaxilla WAS the other maxilla all along. Note while they agreed with my reinterpretation of pedal phalangeal assignments, their materials list incorrectly says supposed IV-2 is
actually III-2 and supposed III-1 is actually II-2, which besides just
being wrong anatomically and contradicting their own figures would leave us with
an extra III-2 and no III-1. They list three
"Withdrawn elements:
left pterygoid; posterior caudal vertebra; left ischium" without
comment, but Hattori (pers. comm. 3-4-2022) indicates the pterygoid and
ischium cannot be identified in the existing materials and that the
caudal is theropodan but not part of the holotypic block. The Lori paper recovered "Fukuivenator" as the most basal alvarezsauroid but found it could be a therizinosaur with just two steps. Hattori et al. actually recovered the latter topology, but scoring the new data in the Lori matrix leaves it sister to <i>Ornitholestes </i>just outside Maniraptoriformes instead.</p><p></p><p><i>Nqwebasaurus </i>had new cranial material described by Radermacher et al. (2021), but they still found it in basal Ornithomimosauria using Choiniere's TWiG matrix. Adding the new data to the Lori analysis resolved it as sister to the alvarezsauroid-therizinosaur clade, but details on the prootic and other elements in the eventual description could change things. <a href="https://www.theropoddatabase.com/Ornithomimosauria.htm#Pelecanimimuspolyodon"><i>Pelecanimimus</i></a>' postcrania were finally described by Cuesta et al. (2022), and incorporating this data moved it to Ornithomimosauria as in the consensus. But it moves back to Alvarezsauroidea in only five steps (which moves <i>Nqwebasaurus </i>back there too), so it's more important than ever to get the skull (and braincase in particular) described.</p><p><i><a href="https://www.theropoddatabase.com/Ornithomimosauria.htm#Khulsanurusmagnificus">Khulsanurus</a> </i>is a recently described alvarezsauroid (Averianov and Lopatin, 2021), but the element described as a distal pubis seems too
small to be this bone, and differs from theropod pubes in the
posteromedial flange and anteroposteriorly deep apron.
Considering its size and shape, this is more likely to be a proximal
dorsal rib. Another new taxon is the parvicursorine <a href="https://www.theropoddatabase.com/Ornithomimosauria.htm#Parvicursorremotus"><span style="font-style: italic;">Ondogurvel alifanovi</span></a> described by Averianov and Loptatin (2022). They reported it differs from <span style="font-style: italic;">Parvicursor</span>
"by the dorsally arcuate supraacetabular crest of the ilium, by the
tibia less curved labially in transverse plane, by less expanded
proximal part of the fibula, and by a relatively shorter pedal phalanx
II-1." While these differences are real, the femur was said to be
"almost identical with the femur in <span style="font-style: italic;">Parvicursor</span>"
and the two emerge as sister taxa if entered into the Lori matrix. As they are from the same formation and <span style="font-style: italic;">Parvicursor</span> is a juvenile with a tibia 62% of the size of <span style="font-style: italic;">Ondogurvel</span>,
it is hypothesized here that the taxa are synonymous and the shorter
pedal phalanx and fused metatarsus are ontogenetic differences while
the others could be individual variation (tibial curvature is known to
vary in <span style="font-style: italic;">Microraptor</span>, for instance). Averianov and Lopatin added <span style="font-style: italic;">Ondogurvel</span> to a TWiG analysis and used implied weighting to recover it sister to <span style="font-style: italic;">Albinykus</span>, but in their earlier <span style="font-style: italic;">Parvicursor</span>
redescription noted this was done because equal weighting led to large
polytomies in Parvicursorinae. Thus it does not strongly argue
against synonymy with <span style="font-style: italic;">Parvicursor</span>. More material of the Bissekty alvarezsaurid were just described by Averianov and Sues (2022 online), which they name <a href="https://www.theropoddatabase.com/Ornithomimosauria.htm#Dzharaonyxeski"><i>Dzharaonyx</i></a>. As with other Bissekty taxa, remains are
isolated so may represent more than one taxon. In this case, each
referred ulna differs in that "The round facet for the aponeurosis
tubercle of the radius is clearly present in the mature specimen ZIN PH
2020/16 but is absent in the juvenile ZIN PH 2021/16", and manual
ungual I examples differ in that "In two specimens (ZIN PH 2445/16 and
2446/16), the collateral grooves are enclosed by notches whereas they
pass through ventral foramina on the third specimen (ZIN PH 2618/16)."
While these could be individual variation, they could also indicate
different taxa.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/a/AVvXsEifMmRo_2coDJpuIy3-FUbbopscnPEz_mU7gZnGTQcIf_uvi0fsscDZ4_8_px5cCkgsuieVv6NqQdKQQFyBSNUpycq5RbmangnX7BeZ8yzcHHyKY3M8jIB35Msbbm3zLAUK3Fu3mhtlF6_TtU6xL-d08ySFcnH32jf7nDgpJzDlIceH1k_l2fjQsK9j=s3072" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="3072" data-original-width="2304" height="320" src="https://blogger.googleusercontent.com/img/a/AVvXsEifMmRo_2coDJpuIy3-FUbbopscnPEz_mU7gZnGTQcIf_uvi0fsscDZ4_8_px5cCkgsuieVv6NqQdKQQFyBSNUpycq5RbmangnX7BeZ8yzcHHyKY3M8jIB35Msbbm3zLAUK3Fu3mhtlF6_TtU6xL-d08ySFcnH32jf7nDgpJzDlIceH1k_l2fjQsK9j=s320" width="240" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Shuvuuia deserti</i> IGM 100/975. Courtesy of the AMNH.<br /></td></tr></tbody></table><br /></p><p></p><p>Eudromaeosaurs have gotten quite a bit of recent attention, with <a href="https://www.theropoddatabase.com/Dromaeosaurs.htm#Vectiraptorgreeni">"Vectiraptor"</a> from England and lots of new data on Mongolian taxa. "Airakoraptor" was officially named <i><a href="https://www.theropoddatabase.com/Dromaeosaurs.htm#Airakoraptor">Kuru</a> </i>(Napoli et al., 2021; Ruebenstahl et al., 2021), and again I express my displeasure for giving any taxon a genus name that is already a word because it's terrible for finding mentions in the literature or online. Just call it "Kururaptor"- that word didn't exist before, so it would be very easy to search for every reference to it. The <i>Kuru </i>talk and paper also clarified what's been a confusing issue, that supposed <i>Velociraptor </i>ulna IGM 100/981 was called that due to a clerical error, and is actually part of partial skeleton IGM 100/3503. Although called <i>Velociraptor </i>sp. by those works, I would say it falls within the range of variation of <i>Tsaagan</i>. Another Mongolian eudromaeosaur issue is the new long-snouted species of <i><a href="https://www.theropoddatabase.com/Dromaeosaurs.htm#Velociraptormongoliensis">Velociraptor</a> </i>based on IGM 100/982, which has been hinted at since Kundrat (2004) and will be finalized once Powers' (2020) thesis chapter is turned into a paper. What might not be as obvious is that the specimen is responsible for a good portion of what we thought was described <i>V. mongoliensis</i> anatomy thanks to Norell and Makovicky (1999), especially the manus. This makes description of fighting dinosaurs <i>V. mongoliensis</i> skeleton IGM 100/25 even more important.</p><p></p><p>The new Crato bird pes <i><a href="https://www.theropoddatabase.com/Ornithothoraces.htm#Kaririavismater">Kaririavis</a> </i>was described by Carvalho et al. (2021 online). It's weird in mixing euornithine characters like a plantarily displaced third metatarsal and hypotarsus with primitive characters like no distal fusion and a low intercondylar eminence. The authors added it to O'Connor's bird analysis and stated it emerged in
a large ornithuromorph polytomy with <span style="font-style: italic;">Piscivoravis</span>, <span style="font-style: italic;">Yanornis</span>, <span style="font-style: italic;">Yixianornis</span>, <span style="font-style: italic;">Songlingornis</span>, <span style="font-style: italic;">Iteravis</span>, <span style="font-style: italic;">Gansus</span>,
hongshanornithids, <span style="font-style: italic;">Apsaravis</span>, <span style="font-style: italic;">Ichthyornis</span>, hesperornithines and
neognaths in a reported 15 MPTs of 1247 steps. However their
supplementary info shows "the 15 Most Parsimonious Trees (MPTs)
resulted in present phylogenetic analysis", but in all of these
<span style="font-style: italic;">Kaririavis</span> is the sister taxon to <span style="font-style: italic;">Ichthyornis</span>. Running the matrix
actually results in 1104 shorter trees of 1246 steps where <span style="font-style: italic;">Kaririavis</span>
can go anywhere in Ornithothoraces except Hesperornithes+Aves and Schizoouridae sensu Wang et al.,
including trees where it is an enantiornithine. Just to make it more complicated, Carvalho et
al. claim that "35 characters [were] treated as ordered" when their
included NEXUS file leaves all characters unordered by default, and
when those characters (identified from the <span style="font-style: italic;">Mengciusornis</span> analysis this was taken from) <u>are</u> ordered, 720 MPTs of 1274 steps are found where <span style="font-style: italic;">Kaririavis</span> is the basalmost euornithine in each one. When added to the Lori analysis <span style="font-style: italic;">Kaririavis</span>
can equally easily fall out as an enantiornithine or a neognath, and
with a single extra step can be sister to all other euornithines with a
plantarily displaced third metatarsal. But the matrix isn't great at handling isolated metatarsi, so I'm not too confidant about any specific placement and just keep it as Ornithothoraces incertae sedis for now.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/a/AVvXsEjpX5YbC1IOaiBsmVlys-8Tj74k1KNBox09NnKycB4ocsjKRlHg2_JByopf_5TQVTMOKXV7Ij6bMKbvyFdwa3z9j0Rb2FQJhzkrBYucH6KGKdGcK_5QMI8iQzlHLmS9hWvneZFda8piJimcj7wEeKwhzdJr7vjB6QtxDfKRpXF3uQLLBa7fi_cEfQ1d=s2048" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1536" data-original-width="2048" height="240" src="https://blogger.googleusercontent.com/img/a/AVvXsEjpX5YbC1IOaiBsmVlys-8Tj74k1KNBox09NnKycB4ocsjKRlHg2_JByopf_5TQVTMOKXV7Ij6bMKbvyFdwa3z9j0Rb2FQJhzkrBYucH6KGKdGcK_5QMI8iQzlHLmS9hWvneZFda8piJimcj7wEeKwhzdJr7vjB6QtxDfKRpXF3uQLLBa7fi_cEfQ1d=s320" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Sacrum of <i>Bambiraptor feinbergi</i> holotype AMNH 30556 in ventral view, anterior to left. Courtesy of the AMNH.<br /></td></tr></tbody></table><br /></p><p>Two new enantiornithines were named- <a href="https://www.theropoddatabase.com/Ornithothoraces.htm#Brevirostruavismacrohyoideus">"Brevirostruavis"</a> and <a href="https://www.theropoddatabase.com/Ornithothoraces.htm#Beiguorniskhinganensis"><i>Beiguornis</i></a>. The latter's description sadly lacks a measurements table, but the photos are better quality than usual for this kind of paper. As for "Brevirostruavis", it was presented as a short-beaked bird with long ceratobranchials, but when reconstructed the beak isn't all that short, and no data is ever given as to shorter ceratobranchial length in other Jehol enantiornithines. This seems to be a trend with Jehol enantiornithines, where the splashy headline is a supposed weird characteristic that doesn't end up having great evidence- horned <i>Dapingfangornis</i>, predatory-beaked <i>Boluochia</i>, <i>Cathayornis caudatus</i> with lots of unfused caudal vertebrae, tiny <i>Liaoxiornis</i>, <i>Chiappeavis </i>with a weird large premaxilla, heterodactyl <i>Dalingheornis</i>. etc.. I think that once Jehol enantiornithines are examined statistically with taphonomy and ontogeny in mind (e.g the huge changes in sternal ossification and thus shape throughout growth), a large number of genera are going to be sunk.</p><p>Finally, we have the euornithines of which the new taxa are O'Connor et al.'s (2021, 2021 online) <a href="https://www.theropoddatabase.com/Ornithuromorpha.htm#Meemannavisductrix">"Meemannavis"</a> and <a href="https://www.theropoddatabase.com/Ornithuromorpha.htm#Brevidentaviszhangi">"Brevidentavis"</a>. The former was found in a huge polytomy by O'Connor et al. in equally weighted strict consensus trees (note the
trees in their figure are majority rule and implied weighting), but sister to <i>Archaeorhynchus </i>in the Lori analysis, which seems plausible. I think this recent trend (also seen in the <i>Ondogurvel </i>paper, <i>Parvicursor </i>redescription and just this week the <i>Dzharaonyx </i>paper) to present implied weighting trees as a solution to a big polytomy is misleading. As the new <i>Musivavis </i>paper shows (Wang et al., 2022- Fig. 14), weighting can give different results from the unweighted trees we've been using for decades and different strengths of weighting can give different topologies from each other. The alvarezsaurid papers used a k value of 3, O'Connor et al. used a k value of 12, and none of them even try to justify their choice. Thus instead of giving defensible cladograms, we get trees generated from arbitrarily skewing the data. In any case, "Brevidentavis" was presented at SVP at "Brachydontornis" which also made it into a few parts of the resulting paper. Creisler (DML, 2021) revealed "Brevidentavis" is the intended final name, and the paper will presumably be fixed once it gets out of Early View and the taxa are official.<br /><br /><b>References</b>-
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of <i>Velociraptor mongoliensis</i>.
American Museum Novitates. 3282, 45 pp.<br /><br />Kundrat, 2004. Two morphotypes of the <i>Velociraptor</i> neurocranium. ICVM-7 Abstracts. Journal
of Morphology. 260(3), 305.<br /><br />
Powers, 2020. The evolution of snout shape in eudromaeosaurians and its
ecological significance. Masters thesis, University of Alberta. 437 pp.<br /><br />Averianov and
Lopatin, 2021 online. The second taxon of alvarezsaurid theropod dinosaurs
from the Late Cretaceous Khulsan locality in Gobi Desert, Mongolia.
Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.2000976<br /><br />Carvalho, Agnolin,
Rozadilla, Novas, Ferreira Gomes Andrade and Xavier-Neto, 2021 online.
A new ornithuromorph bird from the Lower Cretaceous of South America.
Journal of Vertebrate Paleontology. e1988623. DOI:
10.1080/02724634.2021.1988623<br /><br />Chapelle, Norell, Ford, Hendrickx, Radermacher, Balanoff, Zanno and
Choiniere, 2021. A CT-based revised description and phylogenetic
analysis of the skull of the basal maniraptoran <span style="font-style: italic;">Ornitholestes hermanni</span> Osborn 1903. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 81.<br /><br />
Hattori, Kawabe, Imai, Shibata, Miyata, Xu and Azuma, 2021. Osteology of <span style="font-style: italic;">Fukuivenator paradoxus</span>:
A bizarre maniraptoran theropod from the Early Cretaceous of Fukui,
Japan. Memoir of the Fukui Prefectural Dinosaur Museum. 20, 1-82.</p><p>
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new
dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central
Mongolia. American Museum Novitates. 3982, 47 pp.<br /><br />
O'Connor, Lamanna, Harris, Hu, Bailleul, Wang and You, 2021. First
avian skulls from the Lower Cretaceous Xiagou Formation, Gansu, China.
The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 196.<br /><br />
O'Connor, Stidham, Harris, Lamanna, Bailleul, Hu, Wang and You, 2021
online. Avian skulls represent a diverse ornithuromorph fauna from the
Lower Cretaceous Xiagou Formation, Gansu Province, China. Journal of
Systematics and Evolution. Early View. DOI: <span class="epub-doi">10.1111/jse.12823</span><br /><br />
Radermacher, Fernandez, de Klerk, Chapelle and Choiniere, 2021.
Synchrotron μCT scanning reveals novel cranial anatomy of the enigmatic
Early Cretaceous South African coelurosaur, <span style="font-style: italic;">Nqwebasaurus thwazi</span>. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 213-214.<br /><br />Ruebenstahl, Napoli, Bhullar, Turner and Norell, 2021. Two new
eudromaeosaurs from Khulsan (central Mongolia) reveal modern-like
faunal predatory structure amoung non-avian dinosaurs. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 222-223.<br /><br />
Averianov and Lopatin, 2022 online. A new alvarezsaurid theropod dinosaur
from the Upper Cretaceous of Gobi Desert, Mongolia. Cretaceous
Research. DOI: 10.1016/j.cretres.2022.105168</p><p>Averianov and Sues, 2022 online. New material and diagnosis of a new
taxon of alvarezsaurid (Dinosauria, Theropoda) from the Upper
Cretaceous Bissekty Formation of Uzbekistan. Journal of Vertebrate
Paleontology. Latest Articles. e2036174. DOI:
10.1080/02724634.2021.2036174 <br /></p><p>
Cuesta, Vidal, Ortega, Shibata and Sanz, 2022 (online 2021). <span style="font-style: italic;">Pelecanimimus</span>
(Theropoda: Ornithomimosauria) postcranial anatomy and the evolution of
the specialized manus in Ornithomimosaurs and sternum in
maniraptoriforms. Zoological Journal of the Linnean Society. 194(2),
553-591.</p><p>Wang, Cau, Luo, Kundrat, Wu, Ju, Guo, Liu and Ji, 2022. A new
bohaiornithid-like bird from the Lower Cretaceous of China fills a gap
in enantiornithine disparity. Journal of Paleontology. FirstView. DOI:
10.1017/jpa.2022.12 <br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com6tag:blogger.com,1999:blog-3248412803814730250.post-19144679366751731392021-11-08T05:41:00.001-08:002021-11-08T05:59:37.813-08:00Neimengornis the chimaera, "Yuanchuavis" is another Pengornis and more - October 2021 Database update<p>It's time for another Database update. The most extensive thing I did this month was update all of the links, largely due to the DML Archives being down but also affecting quite a few others.</p><p>The coelophysoid <i>Pendraig </i>was added. If anyone has <a href="https://ethos.bl.uk/OrderDetails.do?uin=uk.bl.ethos.488861 ">Warrener's 1983 thesis</a> originally featuring the material, feel free to send a copy over. I also researched the nomen nudum <i>Sinosaurus </i>"shawanensis" that was previously represented by one DML post by Olshevsky in 2002, based on my hard copy of The Jurassic System of China-</p><p>Olshevsky (DML,
2002) reported that within an incomplete set of pages he had, "On p. 9
and p. 17 the paper notes from the Lufeng Formation the species
Sinosaurus shawanensis (Young) among a number of well-known dinosaur
names." He listed the reference as "Stratigraphy of China,
Jurassic System, Summary, Chinese Academy of Geological Sciences, May
1979." and attributed the name to Anonymous, as there was no indication
of the author in the pages he possessed. While such a publication
has never surfaced, an identical situation is present in Cheng (1985),
a section of "The Jurassic System of China", volume 11 in the
Stratigraphy of China series. Perhaps Olshevsky's paper was a
summary of this series printed prior to the series itself, which began
in 1982. In any case, Cheng lists "<span style="font-style: italic;">Sinosaurus shawanensis</span>
(Young)" alongside other taxa from his layer 5 of the Dark Red Beds of
the Lufeng Formation (equivalent to layer 6 of Luo and Wu, 1994).
Notably this is the only species with the author listed in parentheses,
which would normally indicate a species named <span style="font-style: italic;">shawanensis</span> by Young was transferred to <span style="font-style: italic;">Sinosaurus</span> by someone else, but while Young did name <span style="font-style: italic;">Sinosaurus triassicus</span> neither he nor anyone else named a vertebrate species <span style="font-style: italic;">shawanensis</span> (the only animals with that species name before 1985 are brachiopod <span style="font-style: italic;">Cryptospirifer shawanensis</span> Jing et al., 1974 and small shelly fossil <span style="font-style: italic;">Phyllochites shawanensis</span> Duan, 1983, neither from the Lufeng Formation). Young (1951) did refer one tooth from Shawan to <span style="font-style: italic;">Sinosaurus</span> (IVPP V279), but this was to <span style="font-style: italic;">S. triassicus</span>
and was from the Dull Purplish Beds, so doesn't match stratigraphically
with Cheng's taxon. As the holotype and most paratypes of <span style="font-style: italic;">S. triassicus</span>
are from the Dark Red Beds, I think Olshevsky was correct when he noted
"Perhaps it is significant that Sinosaurus triassicus is not listed,
which might mean that Sinosaurus shawanensis is a synonym", as
indeed <span style="font-style: italic;">S. triassicus</span> is not listed by Cheng either. Thus "shawanensis" is near certainly a typo for <span style="font-style: italic;">triassicus</span>, but is still listed here as this can probably not be proven more than thirty years after the fact with the author dead. <br />
Note Molina-Perez and Larramendi (2019) represent <span style="font-style: italic;">Sinosaurus</span> "shawanensis" with isolated dorsal vertebra IVPP V31, which was referred to <span style="font-style: italic;">Sinosaurus triassicus</span>
by Young (1948). Yet its size and morphology are similar to mid
dorsals of sauropodomorph skeleton IVPP V100, also referred to <span style="font-style: italic;">S. triassicus</span>
by Young (1951) in an example of that era's habit of combining
sauropodomorph postcrania with jaws and teeth of carnivorous
archosaurs. Contra Molina-Perez and Larramendi, it was not "More
recent than <span style="font-style: italic;">Sinosaurus triassicus</span>", being from the Dark Red Beds as well, and has no connection to the name "shawanensis." <br /></p><p>Speaking of Molina-Perez and Larramendi's book, I updated several nomina nuda based on data published in it. It's a pretty impressive tome with excellent illustrations, although they are often of taxa based on fragments and thus almost entirely hypothetical. One aspect of maintaining a list of taxa is where to draw the line between nomina nuda proposed as taxa (informally or incompletely) and nicknames. Prior to the book, I had "Weenyonyx" as a nickname, but given its presence in the book's catalogue of spinosaurids I have now provided it with its own entry. Baryonychine spinosaurids got retooled a bit due to new Wessex taxa <i>Ceratosuchops </i>and <i>Riparovenator</i>. As a lumper my initial hunch was that they were individual variations of <i>Baryonyx walkeri</i>, but Barker et al. (2021) do an impressive job detailing all of the differences and similarities between these and '<i>Suchomimus</i>' (<i>Cristatusaurus </i>on my site). Based on the known anatomical data I don't think we can either synonymize the Wessex taxa or sink them into <i>Baryonyx </i>without also doing so to the Nigerian taxon and thus all baryonychines. Interestingly, both Wessex taxa lack premaxillary crests unlike Cristatusaurus but they differ in the other characters in which <i>Cristatusaurus</i> and <i>Suchomimus</i> resemble each other
more than <i>Baryonyx</i> - a less rounded anterodorsal margin to their
premaxillae (also in <span style="font-style: italic;">Ceratosuchops</span> but not <span style="font-style: italic;">Riparovenator</span>); a comparatively larger second alveolus (also in <span style="font-style: italic;">Riparovenator</span> but not <span style="font-style: italic;">Ceratosuchops</span>).<br /><br />I also finally moved <i>Camarillasaurus </i>to Spinosauridae, which was discovered back in 2019. While Samathi et al. (2021 online) correctly reported the supposed cervical MPG-KPS24 was much too small top belong to the type, they still stated the preserved centrum articular surface was posterior. Yet as the parapophysis is adjacent, this is a concave anterior articular surface. Also, the labeled
parapophyses in Sanchez-Hernandez and Benton's (2014) figure 3B are the neural arch peduncles and the supposed
pneumatic foramen is merely a fossa. The combination of large
hypapophysis, concave to flat anterior articular surface and no dorsal
pleurocoels suggests a basal alvarezsauroid, <span style="font-style: italic;">Mahakala</span> relative, troodontid or pygostylian, so it is here assigned to Maniraptora. The supposed coracoid strongly differs from ceratosaurian or
megalosauroid theropods in being anterodistally expanded as in many
paravians, having a thick lip on the lateral side of the posterior
edge, and having a ventrally located foramen that is most likely
damage. The element is near certainly not a coracoid, although a
proper identification would benefit from figures in multiple
perspectives.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiLQb2CqP9KCzWGAMv6_vfdP11IexZ6xCK27-hiwynTbgHgTyvhHkIqYftscr52cNmOwwyRwKwtipSe155bV7Q6fkjyBLSZ1WlwYVMFUxl7t3idY4jGqiwAGUvyiKRC1QN9f9rbB_ntr5M/s611/Camarillasaurus+maniraptoran+dorsal.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="592" data-original-width="611" height="310" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiLQb2CqP9KCzWGAMv6_vfdP11IexZ6xCK27-hiwynTbgHgTyvhHkIqYftscr52cNmOwwyRwKwtipSe155bV7Q6fkjyBLSZ1WlwYVMFUxl7t3idY4jGqiwAGUvyiKRC1QN9f9rbB_ntr5M/s320/Camarillasaurus+maniraptoran+dorsal.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Maniraptoran anterior dorsal (MPG-KPC24) referred to <i>Camarillasaurus cirugedae</i> by Sanchez-Hernandez and Benton (2014; after Hernandez and Benton, 2014). As noted above, B is anterior view and 'parapophysis' in B is a neural arch peduncle.<br /></td></tr></tbody></table><br /><p>Moving to paravians, I added <i>Papiliovenator </i>and also scored it in the Lori matrix where it emerged sister to <i>Zanabazar</i>. Forcing it to be sister to <span style="font-style: italic;">Linhevenator</span> or <span style="font-style: italic;">Philovenator</span> from the same formation requires four more steps each. I hope it gets a more detailed description in the future, as there are entire portions that are not illustrated and barely or not described by Pei et al. (2021)- manus, coracoid and proximal scapula, pelvis, etc.. </p><p></p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4c_yoWNsD7c55UEGHrw2F2RmTWpGl0Bq6iPYNGeHJtiuY5ioS6Fv2EMw8rB2XTjqg51hLF_GNVj9i-RB82tsPovN8wTVTVsEAphmjjvtkpz7XjM1gGhTcFWyl4NB-mbS5Zoggbu12lJE/s2048/Neimengornis.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="2048" data-original-width="1242" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4c_yoWNsD7c55UEGHrw2F2RmTWpGl0Bq6iPYNGeHJtiuY5ioS6Fv2EMw8rB2XTjqg51hLF_GNVj9i-RB82tsPovN8wTVTVsEAphmjjvtkpz7XjM1gGhTcFWyl4NB-mbS5Zoggbu12lJE/s320/Neimengornis.png" width="194" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype of <i>Neimengornis rectusmim</i> (IMMNH-PV00122) (after Wang et al., 2021). Note the different sizes of elements on each side of the specimen and other characters noted below that make this a chimaera.<br /></td></tr></tbody></table><i></i><p></p><p><i>Neimengornis </i>was described by Wang et
al. (2021) as a new genus of jeholornithiform, but this specimen appears to be
a chimaera assembled from different individuals. The right
humerus, radius and ulna are 11%, 5% and 10% longer respectively than
the left; right metacarpals I and II are 20% and 15% longer while III
is 8% shorter than the left; on the right manus digit II with a larger
distal articulation is given a small ungual while digit III has a large
ungual matched with its more slender phalanx; there is a slender
phalanx in the left manus that doesn't match any of the phalanges in
the right manus; the left ilium is 84% the size of the right one with a
blunter postacetabular process and less projected ischial peduncle; the
right femur is 10% longer than the left, the right tarsometatarsus 13%
longer, and pedal phalanges are mostly different in length between
pedes. Other probable indications are the differently shaped
scapulae; arrangement of carpals in the left manus with the two small
ones positioned alongside metacarpal I at the bases of metacarpals II
and III; left distal astragalocalcaneum smaller than the right and
seemingly disarticulated from the tibia; and retrices appearing as
featureless narrow dark lines extending halfway down the tail.
With that in mind, the element identification listed above is based on
Wang et al.'s interpretation of the skeleton as articulated, so the
phalanges in particular are likely placed incorrectly and the left
?ulna could be a radius or tibiotarsus. The right scapula, humeri
(e.g. the short deltopectoral crest of the diagnosis) and ilia seem
more similar to <span style="font-style: italic;">Sapeornis</span>, although the skull, tail and furcula are <span style="font-style: italic;">Shenzhouraptor</span>-grade, while the sacrum has an expanded but distally unfused last rib as in <span style="font-style: italic;">Confuciusornis</span>
and the right scapulocoracoid is more like confuciusornithiforms in the
apparent fusion and short coracoid. The radii, left ulna,
tarsometatarsi and pubes all seem to be avialan, although the latter
plausibly had their distal end modified to be shorter and
pointed. The right pes may be <span style="font-style: italic;">Sapeornis</span>,
which also has pedal ungual I largest and is similar in
proportions. Although the robust and straight metacarpal III and
small ungual II might suggest the right manus is from a
confuciusornithiform, phalanx I-1 doesn't extend past metacarpal II,
digit II is much smaller and phalanx III-2 is the tiny one (unlike
confuciusornithiforms and the manus in <span style="font-style: italic;">Dalianraptor</span> where III-1 is), pointing to it being an artificial articulation of elements. Thus we have a situation much like <span style="font-style: italic;">Dalianraptor</span>.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEirS8fNT-xd-EnX0ko34HtxJn9Yr-5Z2FJoVOOVdI7WR_pw13coaAVS8Q5tcyKX-d_OSimO40evtpCzYaH9Yjq6TWtqpi1H1LMKN11CabMM46kEOGxdqsSRdalG2DtGH2SyUG6ojB0ddvs/s981/Yuanchuavis+snout.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="613" data-original-width="981" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEirS8fNT-xd-EnX0ko34HtxJn9Yr-5Z2FJoVOOVdI7WR_pw13coaAVS8Q5tcyKX-d_OSimO40evtpCzYaH9Yjq6TWtqpi1H1LMKN11CabMM46kEOGxdqsSRdalG2DtGH2SyUG6ojB0ddvs/s320/Yuanchuavis+snout.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Snout tip of <i>Yuanchuavis kompsosoura</i> (IVPP V27883). Note supposed fifth premaxillary tooth is upside down as shown in medial view (right) with protruding root appearing to be the tip in lateral view (left), so is probably a dentary tooth. Also that the first tooth is displaced posteriorly from its alveolus as shown in medial view (after Wang et al., 2021).<br /></td></tr></tbody></table><p>Wang et al. (2021) described a supposed new taxon of
pengornithid- <span style="font-style: italic;">Yuanchuavis kompsosaura</span> based on IVPP V27883, a
skeleton missing pectoral girdles and forelimbs. Regarding its
validity, the premaxilla is said to have five teeth but the medial view
of the right premaxilla in figure S1B shows the last tooth is upside
down so that the root protrudes ventrally as if it's a crown tip in
lateral view, and so is probably a dentary tooth.
Similarly, the same figure shows the edentulous tip of the bone is
exaggerated by the first tooth being displaced and artificially angled
posteriorly. The actual edentulous portion is only two FABLs
long, similar to <span style="font-style: italic;">Pengornis</span>, IVPP V18632 and <span style="font-style: italic;">Parapengornis</span>
(right premaxilla; first tooth displaced and loose in left
element). The supposedly short anterior and posterior lacrimal
processes are similar to <span style="font-style: italic;">Parapengornis</span>, IVPP V18632 and <span style="font-style: italic;">Pengornis</span>
except that the posterior process of the latter is longer as
interpreted by O'Connor. The large hypapophyses on dorsals one
and two are difficult to evaluate in other <span style="font-style: italic;">Pengornis</span>
as anterior dorsals are usually disarticulated and oriented at odd
angles when exposed at all. Contra the authors, the dorsoventral
depth of the dorsal central fossae (e.g. D5) are similar to IVPP
V18632, <span style="font-style: italic;">Parapengornis</span> and <span style="font-style: italic;">Chiappeavis</span>. Finally, the short anterodorsal pygostyle processes are also present in <span style="font-style: italic;">Parapengornis</span> and <span style="font-style: italic;">Chiappeavis</span> (eroded in <span style="font-style: italic;">Pengornis</span>), while the elongate anteroventral process is also present in <span style="font-style: italic;">Parapengornis</span>, partly exposed in <span style="font-style: italic;">Pengornis</span> and probably hidden under caudal vertebrae in <span style="font-style: italic;">Chiappeavis</span>. Of the variable characters noted above for <span style="font-style: italic;">Parapengornis</span> and <span style="font-style: italic;">Chaippeavis</span>,
the pygostyle/metatarsus ratio is intermediate (66%), metatarsal I is
42% the length of metatarsal II, and anterior cervicals are elongate,
again showing no pattern and no distinct separated groups of
ratios. Thus like other Jiufotang pengornithids, <span style="font-style: italic;">Yuanchuavis</span> is
synonymized with <span style="font-style: italic;">Pengornis</span> here.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhITejKvGf8IAOKdNDYoFyLHTa-sssURWyy248B_sfuJke55IZkcFDX3_boj1l2S7rYCY37xDRTRLD-h3z0QfpiywFiwwHm2NMIb9YT8AWtpTxZDdR6MzGh_CuDvpTWMDEYSCbRKjXX7_o/s1898/Pengornis+tail+comp.png" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="424" data-original-width="1898" height="71" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhITejKvGf8IAOKdNDYoFyLHTa-sssURWyy248B_sfuJke55IZkcFDX3_boj1l2S7rYCY37xDRTRLD-h3z0QfpiywFiwwHm2NMIb9YT8AWtpTxZDdR6MzGh_CuDvpTWMDEYSCbRKjXX7_o/s320/Pengornis+tail+comp.png" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Tail comparison of <i>Pengornis </i>specimens, left from Wang et al. (2021), right scaled to dorsal exposure of pygostyle length from Wang et al. (2021; top) and O'Connor et al. (2016; bottom). Note the difference between the retrical arrangement and lengths is not nearly as great as Wang et al.'s figure would suggest, with the long central pair being known to be dimorphic in enantiornithines and confuciusornithiforms.<br /></td></tr></tbody></table><p>Finally, "Yuornis" was added without any controversy. Interestingly, it was originally reported in an SVP abstract back in 2011 (Lu et al., 2011). Speaking of SVP, I previously said I would comment on it in this entry, but I think it would be more useful next month.<br /></p><p><b>References</b>-
Young, 1948. On two new saurischians from Lufeng, Yunnan.
Bulletin of the Geological Society of China. 28, 75-90.<br /><br />Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica.
C(13), 1-96.<br /><br />Cheng, 1985. The Lufeng-Dafang Subregion. In Wang, Cheng and Wang (eds.).
The Jurassic System of China. Stratigraphy of China. 11, 185-189. <br /><br />Olshevsky, DML 2002. <a href="https://web.archive.org/web/20181215172409/http://dml.cmnh.org/2002Apr/msg00630.html">https://web.archive.org/web/20181215172409/http://dml.cmnh.org/2002Apr/msg00630.html</a></p><p>Lu, Xu, Zhang, Jia and Chang, 2011. A new gobipterygid bird
from the Late Cretaceous Central China and its biogeographic implications. Journal
of Vertebrate Paleontology. Program and Abstracts 2011, 147.</p><p>Sanchez-Hernandez and Benton, 2014 (online 2012). Filling the ceratosaur
gap: A new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica
Polonica. 59(3), 581-600.</p><p>
O'Connor, Wang, Zheng, Hu, Zhang and Zhou, 2016 (online 2015). An enantiornithine with a
fan-shaped tail, and the evolution of the rectricial complex in early birds.
Current Biology. 26(1), 114-119. <br /></p><p>Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288
pp.</p><p>
Barker, Hone, Naish,
Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New
spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the
European origins of Spinosauridae. Scientific Reports. 11:19340.<br /><br />Pei, Qin, Wen, Zhao,
Wang, Liu, Guo, Liu, Ye, Wang, Yin, Dai and Xu, 2021 online. A new
troodontid from the Upper Cretaceous Gobi basin of Inner Mongolia,
China. Cretaceous Research. Journal Pre-proof. DOI:
10.1016/j.cretres.2021.105052 </p><p>
Samathi, Sander and Chanthasit, 2021 online. A spinosaurid from
Thailand (Sao Khua Formation, Early Cretaceous) and a reassessment of
<i>Camarillasaurus cirugedae</i> from the Early Cretaceous of Spain.
Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.1874372</p><p>Wang, Wang, Guo, Kang, Ma and Ju, 2021. A new jeholornithiform
identified from the Early Cretaceous Jiufotang Formation in western
Liaoning. Geological Bulletin of China. 40(9), 1419-1427. </p><p>
Wang, O'Connor, Zhao, Pan, Zheng, Wang and Zhou, 2021. An Early
Cretaceous enantiornithine bird with a pintail. Current Biology. 31, 1-8.</p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com18tag:blogger.com,1999:blog-3248412803814730250.post-31628526185202105912021-10-21T07:05:00.000-07:002021-10-21T07:05:34.921-07:00Identity of the mysterious "Luckyraptor" plus other new birds<p>Last year I noticed an entry for "<i>Luckyraptor </i>[Anonymous] 2007 [nomen nudum]" on Olshevsky's <a href="https://www.polychora.com/">Dinosaur Genera List</a>, but could not find a source for the name and George didn't remember either. Turns out my Facebook contact Shahen remembered that it was <a href="https://web.archive.org/web/20160809151020/http://dml.cmnh.org/2008Jun/msg00339.html">an old DML post</a> by Harris. He only says "_Luckyraptor eastensis_ (it doesn't say anything about gen. or sp. nov., but I ain't heard of it anywhere else and assume it's new here!)" so I went to the source and requested a scan of the entry (Zhang, 2007). Each taxon in Zhang's book gets a fossil photo, often bad restoration, Chinese description and notably poor English translation of said description.<br /></p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCDOLQHMfObaMh_exrQqXPPFYYczVefW7SzNEqBXR7YBOL1mPkUlmG_vhTbW6zcZtN-_gAF6nsyq6-iSEC7O8reHnjEox46wVBYgvSibEU1JCMffq4Ajqk_sVXjywDPbfaOzKizsJhuAs/s1743/Luckyraptor.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1190" data-original-width="1743" height="218" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCDOLQHMfObaMh_exrQqXPPFYYczVefW7SzNEqBXR7YBOL1mPkUlmG_vhTbW6zcZtN-_gAF6nsyq6-iSEC7O8reHnjEox46wVBYgvSibEU1JCMffq4Ajqk_sVXjywDPbfaOzKizsJhuAs/s320/Luckyraptor.jpg" width="320" /></a></div><p>As you can see, "Luckyraptor eastensis" is based on the <i>Jixiangornis orientalis</i> holotype, and indeed the latter's name translates to auspicious bird from the East. So this was just a translation error that I would hesitate to even call a nomen nudum except the -raptor- versus -ornis difference is genuine and the popularity of Olshevsky's list has led to it to be listed across the internet.</p><p> </p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhwAxMptluvDesK2ampAwcKPTpr47MV4MYY0aPwf2NlL2Xlsz-tGM-sK54Heb7wRuyi4kL0zhlcF7KdEC3FPp7SaO5_8PAj1StGKkv2uw3lo2wiUowmB6M4u4xIZa3I1iIQq2YZIfxT834/s1679/Sinornithosaurus+zhaoi.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1164" data-original-width="1679" height="222" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhwAxMptluvDesK2ampAwcKPTpr47MV4MYY0aPwf2NlL2Xlsz-tGM-sK54Heb7wRuyi4kL0zhlcF7KdEC3FPp7SaO5_8PAj1StGKkv2uw3lo2wiUowmB6M4u4xIZa3I1iIQq2YZIfxT834/s320/Sinornithosaurus+zhaoi.jpg" width="320" /></a></div><p></p><p>Harris mentioned three other theropod nomina nuda in Zhang's book, one of which is <i>Sinornithosaurus </i>"zhaoi". Turns out that's based on QM V1002 that was later briefly described as a <i>Microraptor gui</i> specimen by Xing et al. (2013). I translate Zhang's Chinese text as-</p><p>
</p><p style="margin-bottom: 0in;"><span lang="en"><i>"Sinornithosaurus
zhaoi</i></span><span lang="en"> (New Species) </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en"> Theropod dinosaurs
with more developed feathers. The skull is of moderate height, the
neck is not very long, and the front jaw and beak are short. The
teeth are not as sharp as </span><span lang="en"><i>Sinornithosaurus
millenii</i></span><span lang="en">, but they are thick and not
degenerated. The forelimbs are correspondingly longer. The hind limbs
are longer than the forelimbs, with three curved claws, and the body
is feathered. The tail is very stiff and powerful. It is 45 cm long,
three times the length of the dorsal vertebrae, and has a slender
shaft. It is a type of theropod that feeds on small mammals. The
species name is dedicated to the Chinese dinosaurologist Mr. Zhao
Xijin. </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en">Place of Origin:
Beipiao, Liaoning </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en">Age: Late Jurassic to
Early Cretaceous"</span></p><p style="margin-bottom: 0in;">However, its humerofemoral ratio (~86%) is actually shorter than the <span style="font-style: italic;">S. millenii</span> holotype (~91%) and <span style="font-style: italic;">Microraptor</span> teeth are typically less elongate than <span style="font-style: italic;">Sinornithosaurus</span>'. The slender ischium, short tibia, elongate pedal phalanx IV-4 and long leg remiges all support assigning QM V1002 to <span style="font-style: italic;">Microraptor</span> instead of <span style="font-style: italic;">Sinornithosaurus</span> and it is here considered a junior synonym of <i>M. zhaoianus</i>. "zhaoi" and the following two taxa are nomina nuda as there is no "explicit fixation of a holotype, or syntypes" (ICZN Article 16.4.1).</p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjyf4BqzbLp7C9uQwM_NcaHTvBs8rWh4i9vzC6ni6lOMe5z3tn_unzDMU92t_DJUyUNyGnPpllurKlup8zA0JNLlrZEcW7D1bH1pXPmphD0f2vJN6IHOJTKa5ev2tEXI2uogkw0nwxGn7s/s1173/Jinzhouornis+delicates.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1173" data-original-width="813" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjyf4BqzbLp7C9uQwM_NcaHTvBs8rWh4i9vzC6ni6lOMe5z3tn_unzDMU92t_DJUyUNyGnPpllurKlup8zA0JNLlrZEcW7D1bH1pXPmphD0f2vJN6IHOJTKa5ev2tEXI2uogkw0nwxGn7s/s320/Jinzhouornis+delicates.jpg" width="222" /></a></div><p style="margin-bottom: 0in;">Next we have <i>"Jinzhouornis"</i> "delicates". My translation is-<br /><br />"<i><span lang="en">J</span></i><span lang="en"><i>inzhouornis
delicates</i></span><span lang="en"> (New Species) </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en"> The primitive bird
whose individual is smaller than the petite </span><span lang="en"><i>Liaoxiornis</i></span><span lang="en">,
belongs to the Enantiornithes group. The head is short and round, the
skull is relatively developed, and the jaw has teeth. The outer body
is feathered, and the forelimbs have become wings. There are 3 curved
finger claws, which are relatively developed, the sternal keel is
prominent, the caudal vertebra have fused, and the tail is short.
There are two <i>Lycoptera </i>fossils on the slab of </span><span lang="en"><i>Jinzhouornis
delicates</i></span><span lang="en">. There is a plant fossil in the
middle. This fossil is the smallest bird fossil discovered in the
Late Jurassic-Early Cretaceous Yixian Formation of western Liaoning,
China, and it remains to be studied. </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en">Place of Origin: Yixian
County, Liaoning </span>
</p>
<p style="margin-bottom: 0in;"><span lang="en">Age: Early Cretaceous"</span></p><p style="margin-bottom: 0in;"></p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiayUMyN2-p5ZgBznuHFAOisVCfOpqnZky3xQZK2N4mQKmN4b20dyBfe5qGbbEr1wd_uS_uwwe0WU1GZTQgZt1BISrgoRy0GHrXKOyrZNkXlwXpxv1r2BJJ2lQyHd10dqRqx7z3WRTfbUk/s902/Jinzhouornis+delicates+close.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="902" data-original-width="562" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiayUMyN2-p5ZgBznuHFAOisVCfOpqnZky3xQZK2N4mQKmN4b20dyBfe5qGbbEr1wd_uS_uwwe0WU1GZTQgZt1BISrgoRy0GHrXKOyrZNkXlwXpxv1r2BJJ2lQyHd10dqRqx7z3WRTfbUk/s320/Jinzhouornis+delicates+close.jpg" width="199" /></a></div><span lang="en"></span>
<p></p>
<p>The generic assignment is odd because <i>Jinzhouornis </i>is a confuciusornithid (and indeed, a junior synonym of <i>Confuciusornis</i>), but this is stated to be an enantiornithine. That identification seems to be correct, given the short fused premaxillae preserved left of the main skull, coracoid and humeral shape, tiny diamond-shaped sternum, and manus with short digit I and reduced ungual and seemingly long metacarpal III projecting past II. In addition to the sternal anatomy, the poorly ossified ends of elements and stated small size strongly suggests a young age. The description seems inaccurate in that the round head is probably due to beak elements overlapping the posterior cranium (contra the restoration), the manual unguals are not well developed or necessarily three in number, and the sternum shows no sign of a keel. The other details are congruent with a juvenile enantiornithine, and this specimen is likely indeterminate as are most other juvenile enants (e.g. <i>Liaoxiornis</i>, <i>Dalingheornis</i>, GMV-2158 and 2159).</p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhF8bCQlQ5FVKlY6zKgncJp-jIVCpkihdpGwz4IhN1spM5KLKMznW4pkr23vXHjTECbOU8vEzdIUncuzS0nvhQplqFJWexK7lUCk712ccep_SN2MZhcCehXUfur7WOIHaZtAMllUZHkTmk/s1159/Smallornis.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1159" data-original-width="762" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhF8bCQlQ5FVKlY6zKgncJp-jIVCpkihdpGwz4IhN1spM5KLKMznW4pkr23vXHjTECbOU8vEzdIUncuzS0nvhQplqFJWexK7lUCk712ccep_SN2MZhcCehXUfur7WOIHaZtAMllUZHkTmk/s320/Smallornis.jpg" width="210" /></a></div><p>Finally, there's "Smallornis liaoningica", whose entry I translate as-</p><p>"<span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="0"><span><i>Smallornis liaoningica</i> (new species)</span></span><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="1"><span>
</span></span><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="2"><span> </span></span></span></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="2"><span>The parietal bones are well developed, the posterior edge of the orbit is concave, the eye holes are large, the beak is long, and the fingers and claws are particularly curved.</span></span> <span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="3"><span>The forelimbs are covered with very delicate feathers, and the down feathers on the head are more beautifully decorated. <i>Smallornis liaoningica</i> is a very small arboreal bird.</span></span><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="4"><span>
</span></span><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="5"><span> </span></span></span></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="5"><span>Place of Origin: Yixian County, Liaoning</span></span><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="6"><span>
</span></span><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="7"><span> </span></span></span></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="7"><span>Age: Early Cretaceous"</span></span></span></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="7"><span> </span></span></span></p><div class="separator" style="clear: both; text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqWhK3_09jegoVZsBPC6yZ3LHx5NQtyG4SkelxonJgVrSqfEyaK5aCe_Rg3inwJ-vODWe7GNxTBSRFZ9AiXI-HUQIwm2iicOHfnr5KcqEMUoPp9Il8N9V_z3eW23W2kBADCbF6nMkq1aE/s988/Smallornis+liaoningica+close.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="988" data-original-width="834" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqWhK3_09jegoVZsBPC6yZ3LHx5NQtyG4SkelxonJgVrSqfEyaK5aCe_Rg3inwJ-vODWe7GNxTBSRFZ9AiXI-HUQIwm2iicOHfnr5KcqEMUoPp9Il8N9V_z3eW23W2kBADCbF6nMkq1aE/s320/Smallornis+liaoningica+close.jpg" width="270" /></a></div><p></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="7"><span>This is most of a bird skeleton, only missing one of the hindlimbs below the knee. Unfortunately the available image quality means that it can't be determined whether this is an enantiornithine or euornithine, although the coracoid structure shows it is ornithothoracine. Obviously none of the listed characters are diagnostic past Pennaraptora or so, but I expect enough is preserved to make it valid compared to named Jehol taxa. Until we have a better scan, I'd say leave it as Ornithuromorpha incertae sedis.</span></span></span></p><p><span class="VIiyi" lang="en"><span class="JLqJ4b ChMk0b" data-language-for-alternatives="en" data-language-to-translate-into="zh-CN" data-number-of-phrases="8" data-phrase-index="7"><span>If anyone has further information like specimen numbers, provenance or higher quality scans for "delicates" or "Smallornis", feel free to share.<br /></span></span></span></p><p><b>References</b>-
Zhang, 2007. The Fossils of China. China University of Geosciences Press. 502 pp.</p><p>
Xing, Persons, Bell, Xu, Zhang, Miyashita, Wang and Currie, 2013. Piscivory
in the feathered dinosaur <i>Microraptor</i>. Evolution. 67(8), 2441-2445. <br /></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com11tag:blogger.com,1999:blog-3248412803814730250.post-78302332209330050802021-09-27T19:44:00.001-07:002021-09-27T19:44:54.275-07:00"Megalosaurus" cloacinus and more - September 2021 Database Update<p>Hi everyone. I realize it's been ten months since the last post, and that's because I've been prioritizing updating the Database over writing blogs. As a compromise of sorts and to not force people to constantly check the Database updates page, I decided to try out posting when I update including features that could have made it into their own blog post.</p><p>One thing I've been doing is working my way through Skawiński et al.'s (2017) paper on Polish Triassic dinosaur reports, which in addition to unnamed fragments, also led to the creation of entries for two supposed <i>Megalosaurus </i>species. <i>silesiacus </i>is a generic carnivorous archosauriform tooth too early to be dinosaurian, while <i>cloacinus </i>has been used for basically every carnivorous archosaur tooth from Rhaetian beds of Germany. The interesting thing about the latter is that workers apparently forgot that it was based on lost teeth described by Quenstedt, not the SMNS tooth figured 47 years later by Huene.<br /></p><p><b><i>"Zanclodon" silesiacus </i></b>Jaekel, 1910<br />
= <span style="font-style: italic;">Megalosaurus silesiacus </span>(Jaekel, 1910) Kuhn, 1965<br />
<b>Early Anisian, Middle Triassic<br />
Lower Gogolin Formation, Lower Muschelkalk, Poland<br />Holotype</b>- (University of Griefswalden/Göttinger coll.; lost?) tooth (24x12x5 mm)<br />
<span style="font-weight: bold;">Referred</span>- ?(Geological Museum
of the Polish Geological Institute-National Research Institute coll.)
tooth (Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and
Niedźwiedzki, 2017)<br />
?(Silesian University of Technology, Faculty of Mining and Geology coll.) tooth (37 mm) (Surmik and Brachaniec, 2013)<b><br />
Comments</b>- Jaekel (1910) noted (translated) "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>a
dinosaur tooth from the lower shell limestone of Upper Silesia, which
would probably be the oldest known dinosaur tooth to date. It comes
from the Chorzov strata of the lower shell limestone of Gogolin, Upper
Silesia, and came to me through the kindness of engineer Fedder in
Opole. The crown shown is 24 mm high, 12 mm wide and 5 mm thick, so it
is quite strongly compressed and slightly curved backwards. Its edge is
extremely finely serrated (Fig. 16 A). I call the form, which for the
time being cannot be specified generically, <span style="font-style: italic;">Zanclodon silesiacus</span>. The only difference between [phytosaur <span style="font-style: italic;">Mesorhinosuchus</span>]
and this tooth form lies in the fact that the former is somewhat
thicker, somewhat less bent back, and that no notch can be detected on
the edge." He referred it to Megalosauridae, and Kuhn (1965)
later referred it to the genus <span style="font-style: italic;">Megalosaurus</span>.
Carrano et al. (2012) correctly noted "could be considered as Theropoda
indet., but we cannot rule out the possibility that it represents a
'rauisuchian' archosaur." </span></span></span>Surmik and
Brachaniec (2013) describe a tooth from Gogolin Quarry in which "a poor
state of preservation makes it impossible to identification of the
presence of edge serration, however it still shows a slightly curvature
and specific both sides flattening" and identify it as seemingly
archosaurian. Skawiński et al. (2017) listed this and another
tooth labeled as <span style="font-style: italic;">Megalosaurus silesiacus </span>as other material of <span style="font-style: italic;">Zanclodon silesiacus</span>.
The latter tooth is stated to be serrated mesially and distally with a
density of 12 per 5 mm. They describe the holotype tooth as
"Probably lost" and "lost", and place all three teeth as
Archosauromorpha indet.. They are more specifically referred to
the <span style="font-style: italic;">Teyujagua</span> plus
archosauriform clade here given the recurvature and small serrations,
as authors from Kuhn onward have noted plesiomorphic theropod teeth are
difficult to distinguish from several clades of archosauriforms (e.g.
erythrosuchids, euparkeriids) known from the Anisian. The age is
far too early for <span style="font-style: italic;">Megalosaurus</span> or another neotheropod, and the presence of serrations is unlike <span style="font-style: italic;">Zanclodon</span>,
so neither genus is appropriate. It should also be noted the
three Gogolin teeth differ in shape with the Silesian University
specimen less recurved and less tapered than the other two, while the
Polish Geological Institute specimen is shorter than the holotype and
less concave distally. This could be positional variation, but
given the lack of proposed synapomorphies could easily represent
multiple taxa.<b><br />
References</b>- Jaekel, 1910. Ueber einen neuen
Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der
Gesellschaft Naturforschender Freunde zu Berlin. 5, 197-229.<br />
Kuhn, 1965. Fossilium Catalogus 1: Animalia. Pars 109: Saurischia. Ysel Press. 94 pp.<br />
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.<br />
Surmik and Brachaniec, 2013. The large superpredators' teeth from
Middle Triassic of Poland. Contemporary Trends in Geoscience. 2, 91-94.<br />
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and
Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical
'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical
Biology. 29(4), 442-472.</p><p> <table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg4Md1TRkBB4YmoRmRM_IEorJJHPquhr-zeTClMIGUwuOeHRXqs6UT7VH8D-BJWF_o68UVnxTQavnEdyAgG5REPQ3LItcqIAOJvKKKbLLvs15NNrIbyYP3RTL2nUNSnhsr0Ms4NAGNDpAI/s645/Zanclodon+silesiacus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="645" data-original-width="309" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg4Md1TRkBB4YmoRmRM_IEorJJHPquhr-zeTClMIGUwuOeHRXqs6UT7VH8D-BJWF_o68UVnxTQavnEdyAgG5REPQ3LItcqIAOJvKKKbLLvs15NNrIbyYP3RTL2nUNSnhsr0Ms4NAGNDpAI/s320/Zanclodon+silesiacus.jpg" width="153" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Holotype tooth of <i>"Zanclodon" silesiacus</i> (University of Griefswalden/Göttinger coll.; lost?) in labial (A), basal section (B) and more apical section (C) (after Jaekel, 1910).<br /></td></tr></tbody></table><br /></p><p></p><p><b><i>"Megalosaurus" cloacinus</i></b>
Quenstedt, 1858<br />
= <i>Plateosaurus cloacinus</i> (Quenstedt, 1858) Huene, 1905<br />
= <i>Gresslyosaurus cloacinus</i> (Quenstedt, 1858) Huene, 1932<br />
= <span style="font-style: italic;">Pachysaurus cloacinus</span> (Quenstedt, 1858) Huene, 1932<br />
<b>Rhaetian, Late Triassic<br />Exter Formation, Germany<br />
Syntypes</b>- (lost) two teeth<br />
<b>Referred</b>- ?(GPIT and SMNS coll.) many teeth (Huene, 1905)<br />
?(SMNS 52457) tooth (~25x11x? mm) (Huene, 1905)<br />
?(SMNS coll.) teeth (Roemer, 1870)<br />
? seven teeth (Miller Endlich, 1870)<br />
<span style="font-weight: bold;">Norian-Rhaetian?, Late Triassic</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">'Lisów Breccia', Poland</span><br />
<span style="font-weight: bold;"></span>?(University of Wroclaw coll.; lost) two teeth (Roemer, 1870) <br />
<span style="font-weight: bold;">Early Hettangian, Early Jurassic</span><br style="font-weight: bold;" />
<span style="font-weight: bold;">Calcaire de Valognes, Manche, France</span><br />
?(University of Caen coll.; destroyed) tooth (Rioult, 1978)<br />
<span style="font-weight: bold;">Comments</span>- Quenstedt (1858)
originally described (translated) "barb-shaped teeth, which are sharp
and finely serrated on the concave side, but rounded and smooth on the
convex side" with a large mesioapically placed wear facet that makes
that edge look straight in side view. He also figures a smaller
tooth which has mesial serrations apically that transition to a
rounded edge basally. These teeth do not share any obvious
synapomorphies and differ in elongation (height/FABL ~300% vs. 138%)
and transverse thickness (42% vs. 75% of FABL), so may not belong to
the same taxon. Miller Endlich (1870) figured seven teeth from
the type locality, stating (translated) they "are mostly flat teeth,
slightly curved on one side, with fine serrations on the sharp inner
edge. The convex side, the back, does not seem to be serrated, but it
is not certain." The figured teeth show a wide range of
variation, with figure 13 in particular being stout and unrecurved with
large serrations, similar to the <span style="font-style: italic;">Lucianosaurus</span> paratype and similarly
referrable to Archosauromorpha incertae sedis. The other teeth
have small serrations, with 14 and 18 being straight and 15-17 and 19
being recurved, with 14, 18 and 15 being progressively more
transversely compressed. As with the syntypes, these exhibit
variation which could be positional or interspecific, and share no
obvious characters that connect them to each other or the
syntypes. Roemer (1870) wrote (translated) "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>In
the Stuttgart Museum I saw teeth from the bone breccia of Bebenhausen
near Tubingen, which show the same fine serration of the side edges as
the teeth described by Quenstedt, but are not curved in a sickle shape,
but are straight. It is very likely that these latter teeth belong to
the same dinosaur as the crooked teeth. With these straight teeth from
Bebenhausen, the tooth shown in FIGS. 4 and 5 from the Lisów Breccia
from Lubsza near Woźniki completely coincides. The double-edged tooth,
which is very delicately and regularly notched at the edges, shows a
more strongly curved (outer) and a less curved (inner) surface, both of
which are smooth except for a very fine, irregular wrinkle. There is
also a much smaller tooth of the same type from the same
location." The straight Bebenhausen teeth sound similar to Miller
Endlich's figures 14 and 18, although the illustrated straight tooth
from Lubsza differs from these in having an increased amount of
mesiodistal expansion basally. The Lubsza tooth also has this
marked basal expansion labiolingually, and both types of root expansion
are atypical of dinosaurs, suggesting this is some other type of
vertebrate. Dzik and Sulej (2007) suggested it "may have belonged
to a phytosaur" without evidence but </span></span></span>
Skawiński et al. (2017) stated "phytosaur fossils have not been found
in the upper Keuper strata in Silesia" and instead placed it in
Archosauromorpha indet.. While this could merely mean phytosaurs
were rare in that strata, phytosaur teeth don't seem to have expanded
roots either (e.g. <span style="font-style: italic;">Nicrosaurus</span>), and it could even be a fish
tooth which often have these types of root expansion. Huene
(1905) listed the species as <span style="font-style: italic;">"Plateosaurus" cloacinus</span> within Theropoda,
stating it includes Rhaetian dentary <span style="font-style: italic;">"Zanclodon cambrensis"</span>. In 1908 he places it
in Plateosauridae within Theropoda and states (translated) "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>The
originals can no longer be found. The Tübingen collection still has
several teeth from Bebenhausen and Schloßlesmuehle, which can be
reconciled well with [Quenstedt's] fig. 12 (l. c.), but are larger. The
serrations are coarse and short, the mesial carina does not extend all
the way to the base." </span></span></span>He illustrated a tooth in figure 274 as "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>From
the Rhaetian Bonebed of Bebenhausen near Tübingen. Tooth in nat. Size.
The tip is missing. Original in the natural history cabinet in
Stuttgart." </span></span></span>Regarding <span style="font-style: italic;">cambrensis</span>, Huene states "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>The
teeth have the greatest resemblance to <span style="font-style: italic;">Plateosaurus cloacinus</span> both in
the whole shape and in the serrations. Whether it is really the same or
just a very similar species, of course, cannot be decided with
certainty given the scanty material", which is not explicit enough to
evaluate given published details. </span></span></span>Huene
later (1932) assigns <span style="font-style: italic;">cloacinus</span> to Teratosauridae within Carnosauria,
listed as both <span style="font-style: italic;">Pachysaurus cloacinus</span> (pg. 6) and <span style="font-style: italic;">Gresslyosaurus
cloacinus</span> (pg. 72, 114). Steel (1970) calls it <span style="font-style: italic;">Gresslyosaurus cloacinus</span> within Plateosauridae. Buffetaut et al. (1991) mentions "A tooth referred to <span style="font-style: italic;">Megalosaurus cloacinus</span>
Quenstedt, from the Lower Hettangian of the Calcaire de Valognes at
Valognes (Manche), [which] has been mentioned by Rioult (1978a) as
having been destroyed by an air raid on the University of Caen in
1944." Without additional details, it can only be said that the
timing suggests a neotheropod. Carrano et al. (2012) incorrectly
claimed SMNS 52457, apparently the tooth in Huene's (1908) figure 274,
is "the holotype and only specimen" of <span style="font-style: italic;">cloacinus</span>, when Huene stated it was only one of "<span class="VIiyi" lang="en"><span class="JLqJ4b" data-language-for-alternatives="en" data-language-to-translate-into="de" data-phrase-index="0"><span>Many
teeth ... in the stone quarries of the Schoenbuch (e.g. Bebenhausen,
Schloesslesmuehle), Wuerttemberg; in the university collection in
Tubingen and in the natural history cabinet in Stuttgart", and that
Quenstedt's originals were lost. SMNS 52457 could be made into a
neotype, but this must be done explicitly (ICZN Article 75.3) and so
has not been accomplished yet. Carrano et al. say the specimen
"is a serrated, recurved tooth of the form typical for theropods. It is
mesiodistally slender but does not show any diagnostic features and is
therefore Theropoda indet", but other taxa with similar teeth lived in
the Rhaetian (e.g. crocodylomorphs, <span style="font-style: italic;">Daemonosaurus</span>), so is here placed in Archosauriformes indet.. Note Huene (1905, 1908) used <span style="font-style: italic;">"Plateosaurus"</span> as a placeholder genus because until 1911 <span style="font-style: italic;">Plateosaurus</span> was thought to have carnivorous teeth, and used <span style="font-style: italic;">"Pachysaurus"</span> and <span style="font-style: italic;">"Gressylosaurus"</span>
in 1932 because until the 1980s more robust 'prosauropod' postcrania
were still associated with carnivorous cranial elements, while Huene
viewed megalosaurids as Jurassic carnosaurs. Our modern consensus
suggests a Rhaetian theropod is more likely to be coelophysoid or
dilophosaur-grade than megalosaurian, but the genus is still used here
as a placeholder as <span style="font-style: italic;">Megalosaurus</span> teeth are more similar to <span style="font-style: italic;">cloacinus</span>' syntypes and SMNS 52457 than prosauropod teeth.</span></span></span><br />
<b>References</b>- Quenstedt, 1858. Der Jura. H. Laupp'schen. 842 pp.<br />
Miller Endlich, 1870. Das Bonebed Württembergs. Druck Von Ludwig Friedrich Fues. 30 pp.<br />
Roemer, 1870. Geologie von Oberschlesien. Robert Nischkowsky. 587 pp.<br />
Huene, 1905. Uber die Trias-Dinosaurier Europas. Zeitschrift der Deutschen Geologischen
Gesellschaft. 57, 345-349. <br />
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung
der Ausseuropäischen vorkommnisse. Geologische und
Palaeontologische Abhandlungen. Supplement 1(1), 1-419. <br />
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte.
Monographien zur Geologia und Palaeontologie. 1, 1-362.<br />
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie. Gustav
Fischer Verlag. 1-87. <br />
Rioult, 1978. Inventaire des dinosauriens mésozoïques de Normandie.
Ecosystèmes continentaux mésozoiques de Normandie (Livret-guide). <span>Université de Caen</span>. 26-29.<br />
Buffetaut, Cuny and le Loeuff, 1991. French dinosaurs: The best record in Europe? Modern Geology. 16, 17-42.<br />
Dzik and Sulej, 2007. A review of the early Late Triassic Krasiejów
biota from Silesia, Poland. Palaeontologia Polonica. 64, 1-27.<br />
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300. <br />
Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and
Niedźwiedzki, 2017 (online 2016). A re-evaluation of the historical
'dinosaur' remains from the Middle-Upper Triassic of Poland. Historical
Biology. 29(4), 442-472.</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg39l7CMWMtb4pOKjuhnDO6Onzq9FJhxWtMHOwyaButb49F0UUeNa7PaJiLcu0-DEn5uW_pETK3AxT9iigunKd4tOCrE1j3jxCKlImmPGLHJG7pOrrm59QywOUjUPzmGnIKK44tV4D6pl8/s588/Megalosaurus+cloacinus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="588" data-original-width="284" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg39l7CMWMtb4pOKjuhnDO6Onzq9FJhxWtMHOwyaButb49F0UUeNa7PaJiLcu0-DEn5uW_pETK3AxT9iigunKd4tOCrE1j3jxCKlImmPGLHJG7pOrrm59QywOUjUPzmGnIKK44tV4D6pl8/s320/Megalosaurus+cloacinus.jpg" width="155" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Syntype teeth of <i>"Megalosaurus" cloacinus</i> (lost) in side and sectional views (after Quenstedt, 1858).<br /></td></tr></tbody></table><br /><p>Other named specimens from the paper include the archosauromorph femur <i><a href="https://www.theropoddatabase.com/Non-theropods.htm#Zanclodonantiquus" target="">Zanclodon? "antiquus"</a></i> based on a museum label, and neotheropod fibula <a href="https://www.theropoddatabase.com/Coelophysoidea.htm#Velocipesguerichi"><i>Velocipes</i></a>. The latter was hilariously assigned to "Vertebrata, nomen dubium" by Rauhut and Hungerbuhler (2000), which is an example of a modern classification philosophy I dislike. Nesbitt et al. (2007) is another egregious example, where a small subset of phylogenetically useful features are checked for, then a specimen is just thrown into whatever clade those features can narrow it to indet.. As opposed to then asking what differences exist between members of that clade and checking the specimen for them, like what Ezcurra and Brusatte (2011) did for <i>Camposaurus</i>. I bet if someone did an in depth study, we would find differences between shuvosaurid and coelophysoid dorsals and caudals and thus be able to narrow down a lot of specimens currently sitting in Archosauria indet. for instance.</p><p>The other thing about Skawiński et al.'s <i>Velocipes </i>discussion that didn't fit into its Database entry is that they claim "It is worth noting that there is great variation in shape of proximal end of fibula of <i>Coelophysis bauri</i> in proximal view – some specimens are more rectangular and others more triangular, some bear flat medial surface, while in others it is strongly concave (compare von Huene 1915, fig. 51; Hutchinson 2002, fig. 2c; Spielmann et al. 2007a, fig. 6 H)." But this is comparing Arroyo Seco AMNH 2745 with Padian's unnamed Petrified Forest coelophysoid (Ezcurra et al., 2021) with the Snyder Quarry coelophysoid, that are not necessarily conspecific.</p><p>Another set of specimens I've been adding/updating over the past several months are Triassic American records, with Arizona and the few bits from Utah complete, I'm currently getting through Texas. That included Camp's material such as <i><a href="https://www.theropoddatabase.com/Non-theropods.htm#Spinosuchuscaseanus">Spinosuchus</a> </i>(that is seemingly a <i>Trilophosaurus </i>species), and revising the <a href="https://www.theropoddatabase.com/Coelophysoidea.htm#Protoavistexensis"><i>Protoavis</i></a> entry. There's a lot of new detail in the latter such as the braincase being compatable with what we now know of non-averostran theropods, the skull roof matching <i>Megalancosaurus</i>, Paul's herrerasaurian characters no longer making sense, the coelophysoid proximal femur being robust and thus probably not juvenile, and commentary on the Kirkpatrick Quarry materials.</p><p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhizwp-ZtRe4ogPhwF3zxA7FoSOMT4dK3oT2zOOp27p93JI2UH98kR3dovLFSEcLAJQSvYaeAAomU2DoRnkueM6NEffBcpBiWHyGiu19t39riB8AI6laCVzRtHslHTcka7UvA_9qvaHmnE/s523/Protoavis+femur+comp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="523" data-original-width="482" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhizwp-ZtRe4ogPhwF3zxA7FoSOMT4dK3oT2zOOp27p93JI2UH98kR3dovLFSEcLAJQSvYaeAAomU2DoRnkueM6NEffBcpBiWHyGiu19t39riB8AI6laCVzRtHslHTcka7UvA_9qvaHmnE/s320/Protoavis+femur+comp.jpg" width="295" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;"><i>Protoavis texensis</i> holotype proximal femur (TTU-P9200; top) in anterior view (after Chatterjee, 1991) compared to <i>"Megapnosaurus" kayentakatae</i> holotype (MNA.V.2623; bottom) (after Rowe, 1989). Note the muscle scars in the former typical of robust coelophysoids.<br /></td></tr></tbody></table><br /></p><p>Speaking of coelophysoids, I finally gave up the battle for Bakker's original Neotheropoda concept and changed everything to the current consensus usages of Neotheropoda and Averostra. Similarly, dilophosaurs are now closer to averostrans, but I note that this isn't as well supported as the consensus would have you believe, with Cau's matrix in the <i>Saltriovenator </i>paper changing to coelophysoid <i>Dilophosaurus </i>in only two steps and Nesbitt's matrix as altered by Ezcurra (and partially corrected by myself) needing only three steps. However, Wang et al. (2016) needs a whopping eighteen more steps to place
<span style="font-style: italic;">Dilophosaurus</span> in Coelophysoidea, so that would be interesting to compare to Tykoski's (2005) thesis that took
twenty additional steps to place <i>Dilophosaurus</i> closer to averostrans. As part of the coelophysoid revision, <a href="https://www.theropoddatabase.com/Coelophysoidea.htm#Sarcosauruswoodi"><i>Sarcosaurus</i></a> was updated after Ezcurra et al. (2021).</p><p>New abelisaurid <a href="https://www.theropoddatabase.com/Ceratosauria.htm#Kurupiitaata"><i>Kurupi</i></a> was added. An upcoming goal is to change Abelisauridae to the stem (away from Noasaurus and Ceratosaurus), and probably move Noasauridae outside Neoceratosauria.</p><p>New tetanurines are <a href="https://www.theropoddatabase.com/Carnosauria.htm#Ulughbegsaurusuzbekistanensis"><i>Ulughbegsaurus</i></a> and <a href="https://www.theropoddatabase.com/Tyrannosauroidea.html#Cryptotyrannus">"Cryptotyrannus"</a>, which had previously been referenced as both an ornithomimosaur and tyrannosauroid in the site. I started to sort out and correct the eastern United States records for those groups now that I have the references instead of relying on Ford's list like I did for much of the original Tyrannosauroidea page, but there's more work to be done in that department.</p><p>I also added new unenlagiines <a href="https://www.theropoddatabase.com/Dromaeosaurs.htm#Ypupiaralopai"><i>Ypupiara</i></a> and <a href="https://www.theropoddatabase.com/Dromaeosaurs.htm#Lopasaurus">"Lopasaurus"</a>. I found a quote in Bertini et al. (1993) seemingly referring to the latter in part- "tibia, a femur and metatarsals (all in the
DNPM, Rio de Jineiro) which Price believed included at least two new
genera and families [of coelurosaurs]." Intriguingly they also mention "a partial toothless dentary (with alveoli but lacking teeth) of a coelurosaur from Loc. 99" which sounds like <i>Ypupiara</i>, but unlike the latter is from a different locality than "Lopasaurus".</p><p>Finally, we have new birds that were added to the cladograms but not given entries yet. <i>Youornis </i>is uncontroversial, but <i>Yuanchuavis </i>looks to just be another <i>Pengornis </i>but with typically dimorphic elongate retrices unlike the <i>'Chiappeavis'</i> specimen. <i>Wasaibpanchi </i>is a Malkani taxon that I don't think is identifiable as theropodan or as teeth from available photos, let alone as a valid taxon of enantiornithine. But that requires going over Malkani's new 2021 paper, and the only thing I've incorporated from that so far is changing which sauropod taxa are valid and listing his new sauropod taxonomic groups and definitions.</p><p>See you in a month.<br /></p><p></p><p></p><p><b></b></p><p><b>Additional References- </b>Rowe, 1989. A new species of the theropod dinosaur <i>Syntarsus</i>
from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate
Paleontology. 9(2), 125-136.</p><p>Chatterjee, 1991. Cranial anatomy and relationships of a new Triassic bird from
Texas. Philosophical Transactions of the Royal Society of London Series B. 332(1265),
277-342.</p><p>Bertini, Marshall,
Gayet and Brito, 1993. Vertebrate faunas from the Adamantina and
Marília formations (Upper Bauru Group, Late Cretaceous, Brazil) in
their stratigraphic and paleobiogeographic context. Neues Jahrbuch für
Geologie und Paläontologie Abhandlungen. 188(1), 71-101.</p><p>Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia
15, 75-88.</p><p>Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD
Dissertation. University of Texas at Austin. 553 pp. <br /></p><p>Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243. </p><p>Ezcurra and Brusatte, 2011. Taxonomic and phylogenetic reassessment of the early
neotheropod dinosaur <i>Camposaurus arizonensis</i> from the Late Triassic of
North America. Palaeontology. 54(4), 763-772.</p><p>Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2016. Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148. <br /></p><p>Ezcurra, Marsh, Irmis and Nesbitt, 2021. A revision of coelophysoid
theropod specimens from Petrified Forest National Park, Arizona
(U.S.A.), reveals a new species from the Upper Triassic Chinle
Formation. 34 Jornadas Argentinas de Paleontologia de Vertebrados,
Libro de Resumenes. R16. </p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com18tag:blogger.com,1999:blog-3248412803814730250.post-67739351713982300642020-12-06T17:41:00.000-08:002020-12-06T17:41:40.010-08:00Antarctic Ichthyornis solved<p>So I've been doing some major updates to the Database for what will probably be a New Years upload, including the ornithuromorph section. One rather sad entry as it currently stands is the Antarctic <i>Ichthyornis</i>- </p><p><i><b>I? sp.</b></i> (Zinsmeister, 1985) <br /><b>Late Cretaceous
<br />Seymour Island, Antarctica
<br />Reference</b>- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic Journal of U.S. 20, 41-42. </p><p>Now with Googling I found the <a href="https://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=&ved=2ahUKEwiyoa6H0brtAhXqN30KHfA3B7kQFjAAegQIBxAC&url=https%3A%2F%2Fs3.amazonaws.com%2FAntarctica%2FAJUS%2FAJUSvXXn5%2FAJUSvXXn5p41.pdf&usg=AOvVaw2fGrdBuL9Q_qGS2P_Va2JP">original paper online</a>, which allowed only a bit of improvement- <br /><br /><i><b>I? sp.</b></i> (Zinsmeister, 1985)<br />
<b>Late Maastrictian, Late Cretaceous<br />Lopez de Bertodano Formation, Seymour Island,
Antarctica</b><br />
<b>Material</b>- several elements<b><br />
Comments</b>- Zinsmeister (1985) states "several small bones tentatively identified as belonging to the Cretaceous bird <span style="font-style: italic;">Ichthyornis</span> were discovered in the upper Cretaceous Lopez de Bertodano formation." <br />
<b>Reference</b>- Zinsmeister, 1985. 1985 Seymour Island expedition. Antarctic
Journal of U.S. 20, 41-42.</p><p>So I saw that Zinsmeister worked with Chatterjee in the 80s, who found the <i>Polarornis </i>holotype in the same place two years before that. I emailed Chatterjee about it, who replied-</p><p>"It was misidentified in the field. These were some shark teeth."<br /><br />Mystery solved! But can we do better? Here's an <i>Ichthyornis </i>tooth-</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEidOoqIiDf8T9OI0QSDXmW-IbqCsSTSRU2KSrcmQMPSqnE170EybMTkn2YGMbLLK9H0En_LmX0Sn9PYa20gu7HDrHoKS_Kr2LpMu3E5VylHIT5W4MCArMHwrP-H2gw0voeXK-NMxrKh8ho/s825/Ichthyornis+tooth.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="718" data-original-width="825" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEidOoqIiDf8T9OI0QSDXmW-IbqCsSTSRU2KSrcmQMPSqnE170EybMTkn2YGMbLLK9H0En_LmX0Sn9PYa20gu7HDrHoKS_Kr2LpMu3E5VylHIT5W4MCArMHwrP-H2gw0voeXK-NMxrKh8ho/s320/Ichthyornis+tooth.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Right eleventh dentary tooth of <i>Ichthyornis dispar</i> (YPM 1450) (after Field et al., 2018).</td><td class="tr-caption" style="text-align: center;"><br /></td></tr></tbody></table><p></p><p>And here's the array of shark teeth from the Lopez de Bertodano Formation of Seymour Island (from a January 2011 expedition). Can we find any easily confusable matches?</p><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj0_0CWOJTcyurqhyDZJUt4TCOjKFa5Zakhhqce-xkkmFHvVjSfTXDIdTObQkW0JUFsZ2cbCF1AMRYT_FS1gHFU5mmvMDVkjjgDHH2oULBLCy00zI8XdNbOnWc-euJV4LMPdfEqEGu_r18/s1148/Lopez+de+Bertodano+shark+teeth.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1148" data-original-width="978" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj0_0CWOJTcyurqhyDZJUt4TCOjKFa5Zakhhqce-xkkmFHvVjSfTXDIdTObQkW0JUFsZ2cbCF1AMRYT_FS1gHFU5mmvMDVkjjgDHH2oULBLCy00zI8XdNbOnWc-euJV4LMPdfEqEGu_r18/s320/Lopez+de+Bertodano+shark+teeth.jpg" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Chondrichthyan teeth from the Lopez de Bertodano Formation (scale 10 mm) (after Otero et al., 2014).<br /></td></tr></tbody></table><p></p><p>I think the circled 16 and 17 are pretty decent matches for a field identification, though much larger if compared directly. Figures 6-17 are all identified as Odontaspidae indet., which covers any morphology similar to <i>Ichthyornis</i>. Add in the fact that they were by far the most abundant teeth recovered (8 samples versus 1-3 for the other taxa), and I think we have a nice solution on our hands.</p><p>I wonder how many other weird records are out there that are based on initial misidentification but stay in the literature because nobody ever publishes a correction?</p><p><br /></p><p><b>References</b>- Otero, Gutstein, Vargas, Rubilar-Rogers, Yury-Yañez, Bastías and Ramírez, 2014. New chondrichthyans from the Upper Cretaceous (Campanian-Maastrichtian) of Seymour and James Ross islands, Antarctica. Journal of Paleontology. 88(3), 411-420.<br /><br />Field, Hanson, Burnham, Wilson, Super, Ehret, Ebersole and Bhullar, 2018. Complete <span style="font-style: italic;">Ichthyornis</span> skull illuminates mosaic assembly of the avian head. Nature. 557, 96-100.</p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-861167739044937762020-11-28T00:39:00.002-08:002020-11-28T00:39:54.997-08:00Is Falcatakely a bird?<p>So this week we got the description of the new Maevarano skull <i>Falcatakely forsterae</i> (O'Connor et al., 2020). It's a pseudo-toucan, with a long, tall snout formed mostly by the maxilla unlike modern birds. O'Connor et al. recover it as an enantiornithine using Brusatte et al.'s TWiG analysis and O'Connor's bird analysis. Our problem is that we only have the anteroventral skull preserved, so no braincase, mandible or postcrania. And being a Maastrichtian deposit in Africa, we don't have the most detailed coelurosaur record. Add in the fact beaks are known to evolve fast on islands (as Madagascar was even back then) and we have a potential problem on our hands.</p><p> <table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgHBrNfwcV6B4Ew-RehYWgDmnwuLCeQzefF0B5gezbAdlycSCKwduz-Bl1t2WCjDsK2mJO1-a8SsmTuKLO1QzT3TCbcnR_EVWiTPKfgQteVSRY9W2o1iSOApnLlCELBXXka3JpqXAFZxg0/s1714/Falcatakely+skull.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="675" data-original-width="1714" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgHBrNfwcV6B4Ew-RehYWgDmnwuLCeQzefF0B5gezbAdlycSCKwduz-Bl1t2WCjDsK2mJO1-a8SsmTuKLO1QzT3TCbcnR_EVWiTPKfgQteVSRY9W2o1iSOApnLlCELBXXka3JpqXAFZxg0/s320/Falcatakely+skull.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Follow your nose to the exciting topic of African coelurosaur diversity. Reconstructed holotype skull of <i>Falcatakely forsterae</i> (UA 10015) (after O'Connor et al., 2020).<br /></td></tr></tbody></table><br /></p><p>The Lori analysis places <i>Falcatakely </i>in two potential positions- a therizinosaurid or an omnivoropterygid. The former doesn't make much sense biostratigraphically, but there is a Maevarano synsacrum of the correct size (FMNH PA 741) that was claimed to share characters with <i>Sapeornis </i>by O'Connor and Forster (2010). So there's a possibility. Forcing <i>Falcatakely </i>to be an enantiornithine as in O'Connor et al.'s analyses requires six more steps. Most of the discordant characters relate to the beak, but the wide laterotemporal fenestra would be odd in an enantiornithine. While I was writing this, Andrea Cau published a post on this topic and reported that he recovered <i>Falcatakely </i>as a noasaurid, which would be quite the phylogenetic jump, but it only takes three more steps in the Lori matrix, so is more parsimonious than the enantiornithine option. It falls out as an elaphrosaurine, so could relate to e.g. <i>Afromimus</i>. The non-beak contradictory characters here include a lack of antorbital fossa lacrimal foramen, long posterior lacrimal process and triradiate palatine, which seem more convincing to me. Additional evidence against these latter two positions is the absence of small ceratosaur (<i>Masiaksaurus </i>is twice as big) or large enantiornithine (Maevarano elements are much smaller) postcrania. Andrea reported (translated) "<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">It
takes 6 further steps to place it in Coelurosauria, and in that case it
is a basal dromaeosaurid: interesting in that regard to note that </span></span><i><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Rahonavis</span></span></i><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"> , known from the same Formation, has also been hypothesized to be a basal dromaeosaurid. </span><span style="vertical-align: inherit;">Can we rule out that </span></span><i><span style="vertical-align: inherit;"><span style="vertical-align: inherit;">Falcatakely</span></span></i><span style="vertical-align: inherit;"><span style="vertical-align: inherit;"> is the (still unknown) skull of </span></span><i><span style="vertical-align: inherit;"><span style="vertical-align: inherit;">Rahonavis</span></span></i><span style="vertical-align: inherit;"><span style="vertical-align: inherit;">? </span><span class="" style="vertical-align: inherit;">The estimated dimensions of the two animals coincide." Forcing <i>Falcatakely </i>to be <i>Rahonavis </i>only requires one more step, which is pretty impressive. In their (amazing) osteology, Forster et al. (2020) refer an isolated dentary "found near the <i>Rahonavis </i>holotype (its precise location was not recorded during excavation)" which does not match <i>Falcatakely</i>'s upper jaw, being upcurved and extensively toothed. But it is similar to other unenlagiines like <i>Buitreraptor </i>and <i>Austroraptor</i>. So much as we have two synsacrum types at this size, unenlagiine-like <i>Rahonavis </i>and <i>Sapeornis</i>-like, we have two cranial types, unenlagiine-like and <i>Falcatakely </i>which is <i>Sapeornis</i>-like in the combination of reduced maxillary dentition, triradiate palatine, modified/reduced antorbital fossa, anteriorly limited naris and strong postorbital-jugal articulation.</span></span></p><p><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;"><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto;"><tbody><tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjBW0m3WvvMOgNVwCH2mIT-5hR8CxtqyLEmGPLzd1vNda4ma_G0cOnqMTl-Y3hnbmviByL5o88o7RT7k8Zu6_6Ynpsil1XrTXS7mdmD6dAN7T67SeDQGDL-ohMbRGWuqLc28rWJPPLSmlE/s1068/Rahonavis+dentary.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="351" data-original-width="1068" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjBW0m3WvvMOgNVwCH2mIT-5hR8CxtqyLEmGPLzd1vNda4ma_G0cOnqMTl-Y3hnbmviByL5o88o7RT7k8Zu6_6Ynpsil1XrTXS7mdmD6dAN7T67SeDQGDL-ohMbRGWuqLc28rWJPPLSmlE/s320/Rahonavis+dentary.jpg" width="320" /></a></td></tr><tr><td class="tr-caption" style="text-align: center;">Referred dentary of <i>Rahonavis ostromi</i> (FMNH PA 740) as a transparent CT reconstruction (after Forster et al., 2020).<br /></td></tr></tbody></table><br /></span></span></p><p><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;">Thus my best guess is that <i>Falcatakely </i>is a basal avialan belonging to the same taxon as FMNH PA 741. But this comes with a huge chunk of salt as it is so far removed temporally and geographically from potential comparable sister taxa. Which is actually a common problem with this part of the tree, as shown by <i>Balaur </i>(= <i>Elopteryx</i>?), <i>Hesperonychus</i>, <i>Imperobator </i>and even <i>Rahonavis </i>itself. We compare these Late Cretaceous taxa to our far more complete Early Cretaceous Jehol record and say <i>Hesperonychus </i>is sorta like <i>Microraptor</i>, <i>Falcatakely </i>is kind of like <i>Sapeornis </i>and <i>Balaur </i>is <i>Jeholornis</i>-grade, but North America, Africa and Europe had their own avialan fauna for 70 million years before them that we're basically unaware of. If the only alvarezsauroid we had was <i>Mononykus</i>' holotype, could we place it correctly as a basal maniraptoran? If the only oviraptorosaur we had was <i>Citipati</i>'s skull, would we recover that correctly as the sister taxon of Paraves? I think that's the position we find ourselves in with <i>Falcatakely</i>, and that future discoveries of African small theropods will lead to new interpretations.</span></span></p><p><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;"> </span></span></p><p><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;"><b>References</b>- </span></span>O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from
the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4),
1178-1201.<br /><br />Cau, 2020 online. <a href="http://theropoda.blogspot.com/2020/11/falcatakely-eterodossia-e-pluralismo.html">theropoda.blogspot.com/2020/11/falcatakely-eterodossia-e-pluralismo.html</a><br /><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;"><br />Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian <span style="font-style: italic;">Rahonavis ostromi</span> from Madagascar. Palaeontologia Electronica. 23(2):a31.</span></span></p><p><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;">O'Connor, Turner, Groenke, Felice, Rogers, Krause and Rahantarisoa, 2020. Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature. DOI: </span></span><span style="vertical-align: inherit;"><span class="" style="vertical-align: inherit;"><span class="c-bibliographic-information__value">10.1038/s41586-020-2945-x</span></span></span></p>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-17361919037253922152020-07-24T17:30:00.001-07:002020-07-25T02:59:48.690-07:00The Arguable Identity of ParaxenisaurusHi everyone. In light of Cau's recent post on the supposed new Mexican deinocheirid "Paraxenisaurus normalensis" (Serrano-Brañas et al., 2020), I figured I'd check the taxon out to see what I thought.<br />
<br />
The first thing you might note are the quotation marks surrounding its name, as this is yet another example of authors not including an lsid or reference to ZooBank in their electronic descriptions. ICZN Article
8.5.3. states names published electronically must "be registered in the
Official Register of Zoological Nomenclature (ZooBank) (see Article
78.2.4) and contain evidence in the work itself that such registration
has occurred", and the pre-print is said to be in preparation for Volume 101 of Journal of South American Earth Sciences, cited as August 2020. Thus it gets to join the ranks of "Thanos" and "Trierarchuncus"as theropods that will eventually be validly named this year. But at least it's not stuck in the purgatory of twelve Scientific Reports Mesozoic theropods, which will never be physically published and thus will remain invalid unless outside action is taken.<br />
<br />
One of the big takeaways from Cau's blogpost is that "<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">I
am doubtful about the possibility of referring these elements [the paratypes] to the
same species of the holotype, since there are very few superimposable
elements among the three specimens. Therefore, there is a risk that
</span></span><i><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Paraxenisaurus</span></span></i><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"> , - understood as the sum of all three specimens - is a chimera." After reading the paper, Andrea REALLY undersold this critique. Here are the specimen materials lists, with the overlapping elements highlighted in matching colors-</span></span><br />
<br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">(BENC 2/2-001; proposed holotype) proximal manual phalanx II-2 or
III-3, partial astragalocalcaneum, partial metatarsal II, phalanx II-1
(115 mm), proximal phalanx II-2, partial metatarsal III, proximal
phalanx III-3, <span style="color: blue;">distal metatarsal IV</span>, phalanx IV-1 (104 mm), phalanx
IV-3 (67 mm), phalanx IV-4 (45 mm), <span style="color: red;">partial pedal ungual IV</span><br />
(BENC 1/2-0054) distal metacarpal I, proximal phalanx I-1, partial manual ungual I, distal metacarpal II, distal phalanx II-2<br />
(BENC 1/2-0091) several proximal caudal central fragments (66, 75, 76
mm), proximal metacarpal II, partial metacarpal III, distal femur (155
mm trans), <span style="color: blue;">distal metatarsal IV</span><br />
(BENC 1/2-0092) several distal caudal vertebrae (70, 71 mm)<br />
(BENC 30/2-001) <span style="color: red;">pedal ungual II, pedal ungual III</span></span></span><br />
<br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">As you can see, there's only one strict overlap, with BENC 1/2-0091 sharing a distal metatarsal IV with the proposed holotype, found ~14 kilometers away. The paper lists no proposed apomorphies or unique combination of characters for distal metatarsal IV, and indeed the description states they preserve largely non-overlapping portions-</span></span><br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">"In the holotype, the distal articular surface is fragmented (Figures 11a1 and 11a2); but in the referred specimen (BENC ½-0091), this surface is nicely preserved and has a non-ginglymoid outline (Figures 11b1 and 11b2). The medial condyle is mostly preserved in the holotype (Figure 11a3), but in the referred specimen it is completely broken (Figure 11b3). Conversely, the lateral condyle is broken in the holotype (Figure 11a4), but is well preserved in the referred specimen (Figure 11b4). Collateral ligament fossae are well developed on both condyles and have approximately the same size and depth (Figures 11a3 and 11b4). In cross-section, the shaft of metatarsal IV near the distal end is thicker dorsoventrally than wide."</span></span><br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Needless to say, metatarsal IV has a shaft which is deeper than wide in all ornithomimosaurs, and the preserved ligament fossae are on opposite sides in each specimen (medial in proposed holotype, lateral in 1/2-0091). Below is a figure comparing the two Mexican specimens with <i>Ornithomimus velox,</i> with 1/2-0091 flipped so that all are comparable as left elements. I don't see anything the "Paraxenisaurus" specimens have in common that could diagnose a taxon.</span></span><br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh7wlV-MasrW4OV4H15AYUAlCg8uhyphenhyphen8Yy3ToNdVNfUvuTdgykveT5aJ2EMV1ZJZd6JC93QIx2rYz4Avo1kRXgbxKarNbjXK1NhPKJb7Qbngx94TShs9xMNHvYQ9MvbEyqwFnL28ZFBz9Mo/s1600/Paraxenisaurus+comp.jpg" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="945" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh7wlV-MasrW4OV4H15AYUAlCg8uhyphenhyphen8Yy3ToNdVNfUvuTdgykveT5aJ2EMV1ZJZd6JC93QIx2rYz4Avo1kRXgbxKarNbjXK1NhPKJb7Qbngx94TShs9xMNHvYQ9MvbEyqwFnL28ZFBz9Mo/s320/Paraxenisaurus+comp.jpg" width="188" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Left distal metatarsal IV of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001, intended "Paraxenisaurus normalensis" paratype BENC 1/2-0091 (right element flipped), and <i>Ornithomomus velox</i> holotype YPM 542 in (top to bottom) dorsal, lateral, ventral and medial views ("Paraxenisaurus" after Serrano-<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Brañas</span></span> et al., 2020; <i>Ornithomimus </i>after Claessens and Loewen, 2016).</td></tr>
</tbody></table>
<br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">While no other elements are exactly matched, referred specimen BENC 30/2-001 does include pedal unguals II and III, while the intended holotype has pedal ungual IV. These are again from different localities, although closer this time (~2.8 km), and this time we have characters listed in the diagnosis- </span></span><br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">"(9) distinctively broad and ventrally curved pedal unguals that angled downward with respect to the proximal articular surface and depending on the digit, the proximodorsal process becomes slightly enlarge and changes its position from nearly horizontal to mostly vertical, adopting a lipshaped appearance; and (10) pedal unguals with a rounded, large foramen on the medial side* and a deep ventral fossa that surrounds a strongly developed, ridge-like flexor tubercle."</span></span><br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi-1Ou970uaezLpH0_x-5NlgNmlw_Fnl0pK0kPcnZgnTzoDL0mbZ9qZh2ZVw7YhS7mmtjrxNtN4h4qTtNVq_M6H3T1fbe0zOrdzMTgzqBUGjQ_s12jaoY_wS5xu68ApmH3b3VngWaLie5U/s1600/Paraxenisaurus+pedal+unguals.jpg" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1227" data-original-width="1031" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi-1Ou970uaezLpH0_x-5NlgNmlw_Fnl0pK0kPcnZgnTzoDL0mbZ9qZh2ZVw7YhS7mmtjrxNtN4h4qTtNVq_M6H3T1fbe0zOrdzMTgzqBUGjQ_s12jaoY_wS5xu68ApmH3b3VngWaLie5U/s320/Paraxenisaurus+pedal+unguals.jpg" width="268" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Pedal unguals of (left to right) intended "Paraxenisaurus normalensis" paratype BENC30/2-001 right digit II, right digit III and intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 left digit IV in (top to bottom) right, left, proximal and dorsal views (after Serrano-<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Brañas</span></span> et al., 2020). Green lines point to supposedly natural median foramen </td></tr>
</tbody></table>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Ventral curvature is plesiomorphic, the unguals of BENC 30/2-001 are not broader than other ornithomimosaurs', and ventral angling with the proximal end held vertically is common in theropods and present in e.g. <i>Garudimimus </i>and <i>Beishanlong</i>. The proximodorsal process "changing its position" is using a difference between 30/2-001's mostly horizontal processes and the intended holotype's more vertical process as character, which in itself presupposes they are the same taxon. The ventral fossa surrounding a ridge-like flexor tubercle is also present in <i>Harpymimus</i>, <i>Garudimimus</i>, <i>Beishanlong </i>and large Dinosaur Park unguals (NMC 1349, RTMP 1967.19.145) and is not shown in the intended holotype but is claimed to be "partially broken." This leaves the medial foramen, which might be a valid character in unguals III and IV (II is damaged in that area), but might also be taphonomic, as there are many other small circular areas of damage (e.g. center of proximal surface of ungual IV). While the two unguals in 30/2-001 are similar to each other, that of the intended holotype is more strongly curved, has that smaller more dorsally angled proximodorsal process, is wider in proximal view, and lacks the expanded ventral half characteristic of ornithomimosaurs that is present in the other specimen. But even if these two pedal unguals are correctly referred, they are all that's present in specimen BENC 30/2-001. So they get us nowhere in determining caudal, manual (besides </span></span><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">proximal manual phalanx II-2 or
III-3) </span></span>or femoral morphology.</span></span><br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">The final issue I noticed was the emphasis on "Paraxenisaurus" having a first pedal digit. This would ironically be unlike <i>Deinocheirus</i>, but plesiomorphically shared with <i>Nedcolbertia</i>, "Grusimimus", <i>Garudimimus</i>, <i>Beishanlong</i>, <i>Archaeornithomimus </i>and <i>Sinornithomimus</i>. The character state is based on metatarsal II, where "a facet on the posterior surface of the distal quarter of this shaft, indicates the presence of an articulation area for metatarsal I." The figure shows a longitudinal groove extending down the posterior center of distal metatarsal II, which as anyone who has scored taxa for Clarke's bird matrix could tell you, is not how non-birds attach their hallux to the metatarsus. Hattori (2016) for instance writes in <i>Allosaurus </i>"there is no attachment scar corresponding to the metatarsal I fossa on either medial or plantar aspect of MT II" and in <i>Citipati </i>"there is no obvious attachment scar of MT I on either medial or plantar aspect of MT II." Serrano-</span></span><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Brañas et al. state "in <i>Garudimimus brevipes</i> ... the attachment site is also placed in the same area as in <i>Paraxenisaurus normalensis</i>", but the feature in <i>Garudimimus </i>is a raised scar with sharp medial demarcation from the shaft. As Middleton (2003) recognized, this scar is for the m. gastrocnemius, specifically the m. gastrocnemius pars medialis (Carrano and Hutchinson, 2002), and I'll note it's present even in <i>Gallimimus </i>which lacks pedal digit I (Osmolska et al., 1972: Plate XLIX Fig. 1b). "Paraxenisaurus"'s groove is then more likely to be the m. flexor digitorum longus II tendon, which "passed through the ventral groove in its respective metatarsal to insert serially on each of the pedal phalanges" in e.g. <i>Tyrannosaurus</i> (Carrano and Hutchinson, 2002). </span></span><br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgRj7jolzj5ZYo1TxVR0Yd9_IZ6ZB6ohTXTSKdGkcL3aij9tryiG0cGrMPRs4rYMz1svQwyJQRyFFPWGxmPQ_Mfu5LLRCefmIHeFjJoClbRTflgR6LBYfbFkAynvEu-66-mv0QeTvw5KO8/s1600/Paraxenisaurus+mtI+comp.jpg" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="842" data-original-width="1161" height="232" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgRj7jolzj5ZYo1TxVR0Yd9_IZ6ZB6ohTXTSKdGkcL3aij9tryiG0cGrMPRs4rYMz1svQwyJQRyFFPWGxmPQ_Mfu5LLRCefmIHeFjJoClbRTflgR6LBYfbFkAynvEu-66-mv0QeTvw5KO8/s320/Paraxenisaurus+mtI+comp.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Left metatarsal II in ventral view of (left to right) intended "Paraxenisaurus normalensis" holotype BENC 2/2-001 (after Serrano-<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Brañas</span></span> et al., 2020; yellow arrow points to supposed articulation for metatarsal I), <i>Garudimimus brevipes</i> holotype IGM 100/13 (after Kobayashi and Barsbold, 2005; line points to supposed articulation for metatarsal I), <i>Gallimimus bullatus</i> ZPAL MgD-I/94 (after Osmolska et al., 1972), and <i>Tyrannosaurus rex</i> FMNH PR2081 (after Brochu, 2003).</td></tr>
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<br />
<div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">What exactly is "Paraxenisaurus"? Comparison is hindered by the specimens being figured mixed together, and the figures are not in numerical order in the preprint, being shown in the order of- 1, 10-19, 2, 20-23, 3-9. In addition, the scale bars vary within the same figure (e.g. phalanx IV-1 is proximally ~61 mm wide in figure 14a but ~93 mm wide in figure 14e) and the listed measurements are different yet (e.g. IV-1 is listed as 83 mm wide). Thus any composite reconstruction is necessarily approximate. The supposed manual element is too fragmentary to give much information, but it is of the appropriate size and shape to be a proximal pedal phalanx I-1. This would make more sense preservationally since the other material preserved in the specimen is all from the tarsus and pes. It's a shame the astragalocalcaneum is not described better or figured in more views, as the dorsal (= proximal?) perspective has many broken surfaces and edges, so that e.g. the small calcaneum might be preservational. The fused proximal tarsals are like ceratosaurs, deinocheirids (<i>Deinocheirus </i>plus <i>Hexing</i>), alvarezsaurids and caenagnathids. Having any sense of the ascending process morphology could tell us much. Metatarsal II is not obviously deeper than wide, unlike ornithomimosaurs (except <i>Harpymimus</i>; unreported in deinocheirids), but like carcharodontosaurids, therizinosauroids, some oviraptorids and velociraptorines. The proximal outline of metatarsal III would at first glance appear to be the strangest thing about this material, being reconstructed as strictly dorsoventrally oval unlike all(?) other theropods. Tilting it and adding a posterior tapered tip results in a close match to <i>Majungasaurus </i>however (see figure below). If it is an unreduced proximal metatarsal III, tyrannosauroids, most ornithomimosaurs, alvarezsauroids and pennaraptorans would be excluded. Proximal phalanx II-2 lacks a proximoventral heel, so is not from a deinonychosaur. The pedal phalanges are too elongate to be therizinosauroid, and the pedal ungual is too broad. Phalanges are not as dorsoventrally compressed as <i>Mapusaurus</i>, and as noted above they lack the ventrolateral shelves found in ornithomimosaurs. Abelisaurid phalanges seem similar however. I wonder if we have a case like <i>Camarillasaurus</i> or probably <i>Dandakosaurus </i>involving misidentified elements making the specimen seem stranger than it really was, with </span></span><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">so many edges of supposed metatarsal III dotted to indicate incompleteness that it could actually be metatarsal II or IV</span></span>. Certainly nothing connects this specimen with <i>Deinocheirus</i>. As per the numerous errors illustrated by Hartman et al. (2019) nobody should trust Choiniere et al.'s scorings in any case. The Lori matrix recovers "Paraxenisaurus" as a ceratosaur closest to <i>Aucasaurus </i>as far as taxa with well preserved feet are concerned, but also doesn't include characters particular to ceratosaurs and isn't great with pedal characters in general. So I would place the specimen as Neotheropoda incertae sedis (or even indet.) pending a better description of the tarsus and of the real bone surfaces on supposed proximal metatarsal III. </span></span></div>
<div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span></div>
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<span style="vertical-align: inherit;"><table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="height: 347px; margin-left: auto; margin-right: auto; width: 667px;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEghFZ5QTLKwu7uwX9DUNZybBLPF1U1TUhBiy_CGqV_tjcRhB4IEIx-FtyXCBuEIz0qyYzcu7-VRr_WuxPogbh1JPt676YCkeRDjgkIdoBsA44IjGcVA8YYz9HiSUtevI6EPjyfF8qJ7nOw/s2048/Paraxenisaurus+holotype+pes+comp2.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1385" data-original-width="2048" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEghFZ5QTLKwu7uwX9DUNZybBLPF1U1TUhBiy_CGqV_tjcRhB4IEIx-FtyXCBuEIz0qyYzcu7-VRr_WuxPogbh1JPt676YCkeRDjgkIdoBsA44IjGcVA8YYz9HiSUtevI6EPjyfF8qJ7nOw/s320/Paraxenisaurus+holotype+pes+comp2.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><span style="font-size: x-small;">Pes of "Paraxenisaurus normalensis" holotype (center) in dorsal view compared to <i>Majungasaurus crenatissimus</i> comp<span style="font-family: inherit;">osite (left) and <i>Deinocheirus mirificus</i> referred specimen IGM 100/127 (right). Colored proximal view of "Paraxenisaurus" is after <span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Serrano-Brañas et al., with reori<span style="font-size: xx-small;">ented metatarsal III as per my interpretation shown above that. Note "Paraxenisaurus" elements were scaled using their scale bars, whereas scaling to listed meas<span style="font-family: inherit;">urements results in different proportions, so those should be seen as approximate. "Paraxenisaurus" after Serrano-Bra</span></span></span></span></span><span style="font-family: inherit; font-size: xx-small;"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><span style="font-family: inherit;"><span style="vertical-align: inherit;"><span style="font-family: inherit;"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">ñ</span></span></span></span>as </span></span></span></span><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><span style="font-size: xx-small;">et al. (2020), <i>Majungasaurus </i>after Carrano (2007) and <i>Deinocheirus </i>after Le</span>e et al. (2014).</span></span></span></span></span></td></tr>
</tbody></table>
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<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><b>References</b>- </span></span><span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur, <i>Gallimimus
bullatus</i> n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia.
Palaeontologica Polonica. 27, 103-143.</span></span></span><br />
<div>
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Carrano and Hutchinson, 2002. Pelvic and hindlimb musculature of <i>Tyrannosaurus rex</i> (Dinosauria: Theropoda). Journal of Morphology. 253, 207-228.</span></span></span></div>
<div>
<br />
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Brochu, 2003. Osteology of <i>Tyrannosaurus rex</i>: Insights from a nearly
complete skeleton and high-resolution computed tomographic analysis of the skull.
Society of Vertebrate Paleontology Memoir. 7, 138 pp. </span></span></span></div>
<div>
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"></span></span></span></div>
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span></span>
<br />
<div>
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Middleton, 2003. Morphology, evolution, and function of the avian hallux. PhD thesis, Brown University. 147 pp.</span></span></span></div>
<div>
<br />
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Carrano, 2007. The appendicular skeleton of <i>Majungasaurus crenatissimus</i>
(Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In Sampson
and Krause (eds.). <i>Majungasaurus crenatissimus</i> (Theropoda: Abelisauridae)
from the Late Cretaceous of Madagascar. SVP Memoir 8, 164-179.</span></span></span></div>
<div>
<br />
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Lee, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and Tsogtbaatar,
2014. Resolving the long-standing enigmas of a giant ornithomimosaur <i>Deinocheirus
mirificus</i>. Nature. 515, 257-260.</span></span></span></div>
<div>
<br />
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Kobayashi and Barsbold, 2005. Reexamination of a primitive ornithomimosaur,
<i>Garudimimus brevipes</i> Barsbold, 1981 (Dinosauria: Theropoda), from the
Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521.</span></span></span></div>
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<div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Claessens and Loewen, 2016 (online 2015). A redescription of <i>Ornithomimus velox</i> Marsh,
1890 (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. 36(1), e1034593.</span></span><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span></div>
<div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span></div>
<div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Hattori, 2016. </span></span><span class="NLM_article-title hlFld-title">Evolution of the hallux in non-avian theropod dinosaurs.</span><span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"> Journal of Vertebrate Paleontology. 36(4), e1116995.</span></span></span></div>
<div>
<span class="NLM_article-title hlFld-title"><span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br /></span></span></span></div>
<div>
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247.</div>
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;"><br />Cau, 2020 online. <a href="http://theropoda.blogspot.com/2020/04/paraxenisaurus-un-deinocheiride.html">http://theropoda.blogspot.com/2020/04/paraxenisaurus-un-deinocheiride.html</a></span></span><br />
<br />
<span style="vertical-align: inherit;"><span class="goog-text-highlight" style="vertical-align: inherit;">Serrano-Brañas,
Espinosa-Chávez, Maccracken, Gutiérrez-Blando, de León-Dávila and
Ventura, 2020. <span style="font-style: italic;">Paraxenisaurus normalensis</span>, a large deinocheirid
ornithomimosaur from the Cerro del Pueblo Formation (Upper Cretaceous),
Coahuila, Mexico. Journal of South American Earth Sciences. 101, 102610.</span></span>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com17tag:blogger.com,1999:blog-3248412803814730250.post-23473926701717179042020-06-13T07:27:00.000-07:002020-06-14T07:01:12.821-07:00The Unecessary Death of SteneosaurusNot a dinosaur, but a new paper on the classic crocodylomorph <i>Steneosaurus </i>exemplifies a troubling trend in recent vertebrate taxonomy. Johnson et al. (2020) reexamine the original material of <i>Steneosaurus</i>, an aquatic croc from the Jurassic of France. It hadn't been seriously looked at since the 1860s, so this is one of my favorite kinds of paleontology papers- restudying a fragmentary old specimen in a modern light. What do they find?<br />
<br />
We first get a detailed recount of its history, with two decades as Cuvier's "tête à museau plus allongé" (= head with a more elongated snout; I have to praise the authors for translating all the French to English, even in our spoiled era of Google Translate it saves time), before it was named <i>Steneosaurus rostro-major</i> by Geoffroy Saint-Hilaire in 1825. Eudes Deslongchamps and son tackled it in the 1860s, where they viewed the specimen as too poorly preserved and so "stated that the taxon to represent the genus <i>Steneosaurus </i>should be either <i>‘Steneosaurus’ megistorhynchus</i> Eudes-Deslongchamps, 1866, or <i>‘Steneosaurus’ edwardsi</i> Eudes-Deslongchamps, 1868c." Ha! You don't get to just take somebody's genus and affix your new species as its type. They were the last to examine the specimen in detail however, making that a pretty bad note to end on.<br />
<br />
Johnson et al. then reexamine the type snout of <i>Steneosaurus</i>, correcting the species name by eliminating the hyphen, officially making it the lectotype, noting Steel had determined the posterior skull to be <i>Metriorhynchus</i>, and illustrating and redescribing the specimen. Excellent work and very well done. After eliminating <i>Mycterosuchus nasutus</i>, <i>'Steneosaurus' leedsi</i>, <i>'S.' heberti</i> and <i>Lemmysuchus</i> and other machimosaurins based on numerous dissimilar characters, the authors come to the contemporaneous <i>'Steneosaurus' edwardsi</i>.<br />
<br />
"As mentioned before, this was a second species that Eudes-Deslongchamps (1867–69) considered identical to <i>S. rostromajor</i>. These two taxa share a combination of features including:<br />
1. A subcircular, moderately interdigitating premaxilla-maxilla suture.<br />
2. Maxillae ornamented with irregular grooves.<br />
3. A shallower mediolateral compression of the posterior maxillae, as opposed to <i>‘S.’ heberti</i> (MNHN.F 1890-13).<br />
4. Horizontally flat posterior premaxilla in lateral view.<br />
5. Deep anterior and mid-maxillary reception pits that gradually become shallower towards the posterior maxilla.<br />
6. Subcircular to circular alveoli that remain relatively the same size throughout the maxilla.<br />
7. Teeth with well-pronounced enamel ridges at the base."<br />
<br />
Well how cool is that? They put in the hard work, found the matching more complete specimens, and now we have <i>Steneosaurus edwardsi</i> as a junior synonym of <i>S. rostromajor</i>, giving us a good look at what <i>Steneosaurus </i>really was after two hundred years.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjmTuUcUMiNvU0t9MYGTENT_z0XC8w1TFg42v7pBlQoLkO6aVScqdE8_kXRtOzezS6-IimSUVdL9TqnXKCG5mmhGNIivn5gycgee1qhDzp8vNLoX7aZHk9WPwnZ97KtyPtF95cO9g3YeMk/s1600/Steneosaurus.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1021" data-original-width="858" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjmTuUcUMiNvU0t9MYGTENT_z0XC8w1TFg42v7pBlQoLkO6aVScqdE8_kXRtOzezS6-IimSUVdL9TqnXKCG5mmhGNIivn5gycgee1qhDzp8vNLoX7aZHk9WPwnZ97KtyPtF95cO9g3YeMk/s320/Steneosaurus.jpg" width="268" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Lectotype of <i>Steneosaurus rostromajor</i> (MNHN.RJN 134c-d) in dorsal (A, B) and ventral (C, D) views. (after Johnson et al., 2020).</td></tr>
</tbody></table>
<br />
But no.<br />
<br />
Johnson et al. immediately say "it is important to note that many of these characters may, in fact, be related to sexual dimorphism, ontogeny and intraspecific variation." True, but that could be said for basically every character supposed to diagnose Mesozoic croc genera, or theropod genera, pterosaur genera, etc.. Unless you have some specific example like 'enamel ridges have been shown to develop with age and both <i>S. rostromajor</i> and <i>S. edwardsi</i> are larger than <i>S? leedsi</i> or <i>S? heberti</i> with weak ridges', then it's just hand-waving. And no, Johnson et al. never develop such an argument for one of those characters, let alone all seven.<br />
<br />
Next, we get "In addition to the sexual dimorphism/ontogeny problem, one of the critical issues about MNHN.RJN 134c-d is that it is poorly preserved." Sure, but you were still able to perform many comparisons. Again, the authors never say any of their seven characters are taphonomic, so it's another objection without substance.<br />
<br />
Yet the worst rationale for rejecting <i>Steneosaurus </i>is "in reality, the name <i>Steneosaurus </i>is extremely impractical. It was used for many metriorhynchid specimens (e.g. <i>‘Steneosaurus’ gracilis</i>, <i>‘Steneosaurus’ palpebrosus</i> and <i>‘Steneosaurus’ manselii</i>) during much of the 19th century, largely in part due to Cuvier’s metriorhynchid skull region (MNHN. RJN 134a-b) being attributed to the teleosauroid rostral section (MNHN.RJN 134c-d). Indeed, the concise, classical definition of <i>‘Steneosaurus’</i> as we interpret it today was not given until the work of both Eudes-Deslongchampses (1868c, 1867–69)"<br />
<br />
Substitute <i>Megalosaurus </i>in there to see how ridiculous it is. That has had over 45 species assigned to it, and was named in the 1820s but didn't have a modern concept associated with it until the 1980s. When Johnson et al. lament that for <i>Steneosaurus </i>"rather than comparing characters outright, comparison is by process of elimination (or the question of ‘what features does this specimen lack?’)", that perfectly describes the <i>Megalosaurus </i>paralectotype dentary.<br />
<br />
"After the Eudes-Deslongchampses’ treatment, what was left was an undiagnostic, chimeric type specimen for <i>S. rostromajor</i> (MNHN.RJN 134) and the genus <i>Steneosaurus </i>was redefined using a new type species that was not accepted by some researchers. In addition, since the Eudes-Deslongchampses, there has been no attempt to rectify this taxonomic nightmare;"<br />
<br />
You just showed it was diagnostic, Steel long ago got rid of the chimaeric portion, Eudes-Deslongchamps' stupid attempts to name new type species have no relevance, and you have done the work to finally rectify this taxonomic nightmare.<br />
<br />
"Due to these three significant factors (uncertainty of variable characters, poor preservation and<br />
unreasonable name), we have concluded that <i>S. rostromajor</i>, and therefore <i>‘Steneosaurus’</i> (MNHN.RJN 134c-d), cannot be confidently assigned to an existing teleosauroid species."<br />
<br />
Nope, you just showed it can be assigned to the same species as <i>S. edwardsi</i>.<br />
<br />
Actually, I correct myself. THIS is the worst rationale for rejecting <i>Steneosaurus</i>- "In addition, MNHN.RJN 134c-d was initially diagnosed based on significant orbital and temporal characteristics (from the metriorhynchid MNHN.RJN 134a-b), along with generic rostral ones. Because the skull material is now known to be from a metriorhynchid, this ‘hybrid type specimen’ factor adds to the doubtful validity of <i>Steneosaurus</i>. According to Article 23.8 of the ICZN Code, ‘a species-group name established for an animal later found to be a hybrid (Art. 17) must not be used as the valid name for either of the parental species (even if it is older than all other available names for them)’ (this also signifies that the species name <i>rostromajor </i>is itself invalid). As such, MNHN.RJN 134c-d serves as an undiagnostic specimen; we, therefore, consider MNHN.RJN 134c-d to be a nomen dubium and, as such, <i>Steneosaurus </i>is treated as an undiagnostic genus."<br />
<br />
If the term "parental species" didn't tip you off, Article 23.8 applies to hybrid individuals (those resulting from different species interbreeding), not type specimens chimaerically combined from multiple species. The Article doesn't even say what Johnson et al. think- it says a name for a hybrid can't be used for either of the species that bred to make it, so that e.g. even if a mule's scientific name was erected prior to that of horse's or ass's, it can't be the name for horse or ass. And indeed even the cited Article 17 says that hybrids and chimaeras can be the basis of valid names- "The availability of a name is not affected even if 17.1. it is found that the original description or name-bearing type specimen(s) relates to more than one taxon, or to parts of animals belonging to more than one taxon; or 17.2. it is applied to a taxon known, or later found, to be of hybrid origin..."<br />
<br />
If Johnson et al.'s interpretation were right, there goes <i>Gojirasaurus</i>, <i>Protoavis</i>, <i>Chuandongocoelurus</i>, <i>Chilantaisaurus</i>, <i>Fukuiraptor</i>, <i>Coelurus</i>, <i>Alectrosaurus</i>, <i>Dakotaraptor</i>, etc..<br />
<br />
Before the big reveal, we have in the Conclusion what can only be described as a lie- "Through character comparison-and-elimination, the only taxon with which MHNH.RJN 134c-d could hypothetically be referred to is <i>‘S.’ edwardsi</i>, but the two do not share any clear autapomorphic characters or a unique combination of characters."<br />
<br />
What are your seven listed characters if not "a unique combination of characters"? Does any other teleosaurid have them? If not, they are unique. In any case, we get the motivation for dumping <i>Steneosaurus </i>twice at the end of the paper-<br />
<br />
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action, which we will highlight in a forthcoming paper (Johnson, 2019)."<br />
<br />
"We believe that establishing teleosauroid taxonomy from the beginning with a series of ‘clean’ type species/specimens, with every nomenclatural act correctly formulated, is the best course of action. This will necessitate a revised teleosauroid taxonomy, in which species previously referred to the genus <i>Steneosaurus </i>are given new generic names. This work will be published by us in a separate contribution, based on the comprehensive teleosauroid phylogenetic analysis in Johnson’s PhD thesis (2019)."<br />
<br />
Basically everything I hate about a current trend in vertebrate paleontology- just throw out old specimens and dishonor their authors who correctly reported what was new at the time to come up with your own names. At least dumping <i>Stegosaurus armatus</i> or designating a neotype for <i>Allosaurus fragilis</i> could be claimed to save time and effort actually analysing the types, if you don't want to do the science to figure out if <i>armatus </i>is actually different from <i>stenops </i>or if <i>fragilis </i>can be distinguished from <i>Saurophaganax</i>. But Johnson et al. already did all the hard work and found <i>Steneosaurus edwardsi</i> was <i>S. rostromajor</i>, they would just rather use Johnson's new genus name for the taxon.<br />
<br />
And their reasons are just grasping at straws. 'Sure we identified these seven charactesrs uniquely shared by <i>Steneosaurus rostromajor</i> and <i>S. edwardsi</i>, but uhh.. could be sexually dimorphic? Or anything could be individual variation. Or ontogenetic? Lots of things turn out to be ontogenetic. Plus it's broken. Sooo broken. Sure we could evaluate characters, but who wants a taxon whose holotype isn't pristine? Plus a lot of people had stupid ideas about <i>Steneosaurus </i>over the past two hundred years. What do us scientists do when we have a complicated situation to resolve that was only partially understood historically? Trash their names and give yourselves credit for new genera.' Thus <i>Steneosaurus </i>gets the eternal identity of "all evidence points to it being <i>Johnsonosaurus edwardsi</i>, but ehhh... we just sort of ignore it now as Teleosauridae indet. and it's forgotten."<br />
<br />
To conclude, <i>Steneosaurus </i>is really outside my wheelhouse. But if Johnson et al.'s philosophy spreads, we're in danger of losing a lot of historical taxonomy and deserved credit to lazy or selfish authors. Just look at <i>Microraptor </i>for example, whose holotype of<i> M. zhaoianus</i> lacks a decent skull. Some decades down the line, what if cranial differences support various Jiufotang species and someone's like 'the postcranial proportions are unique between the <i>M. zhaoianus</i> type and <i>M. hanqingi</i>, but I want a complete type specimen, so <i>Microraptor </i>is an invalid undiagnostic nomen dubium, and instead I propose <i>Mybetterraptorgenus hanqingi</i> and <i>M. gui</i>.' Just hope they don't pull a Wilson and Upchurch and claim '<i>Microraptor </i>is invalid and co-ordinate suprageneric Linnean taxa must likewise be abandoned' and replace Microraptorinae with Mybetterraptorgenusinae.<br />
<br />
<b>References</b>- Johnson, 2019. The taxonomy, systematics and ecomorphological diversity of Teleosauroidea (Crocodylomorpha, Thalattosuchia), and the evaluation of the genus '<i>Steneosaurus'</i>. PhD Thesis, University of Edinburgh. 1062 pp. <br />
<br />
Johnson, Young and Brusatte, 2020. Emptying the wastebasket: A historical and taxonomic revision of the Jurassic crocodylomorph <i>Steneosaurus</i>. Zoological Journal of the Linnean Society. 189(2), 428-448.<br />
<br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com17tag:blogger.com,1999:blog-3248412803814730250.post-45928420091785600132020-05-30T07:04:00.001-07:002020-05-30T07:04:55.007-07:00It's finally January 1, 200n and Phylonyms is published!Ah the PhyloCode, the so-called future of biological nomenclature whose release has always kept on slipping ever more distantly into the future. After 20 years of waiting, we now have Phylonyms: A Companion to the PhyloCode, by de Queiroz et al. (2020), "a turning point in the history of phylogenetic nomenclature" according to its introduction. As the book states "Phylonyms serves as the starting point for phylogenetic nomenclature governed by the PhyloCode. According to the preamble, “This code will take effect on the publication of Phylonyms: A Companion to the PhyloCode, and it is not retroactive.” Thus, names and definitions published here have precedence over any competing names and definitions published either before (or after) the publication of Phylonyms." So for anyone invested in standardized phylogenetic nomenclature, this is it. Nothing better is coming down the pipeline in our lifetimes, so let's see what we're stuck with.<br />
<br />
First of all, it's expensive. You can get an ebook for $222 on Amazon or a hardcover sometime after June 9th for $234. I found an electronic version for $169 plus tax on VitalSource, but you have to use their reader. It's 1323 pages though, so isn't a bad deal. That's less than five $37.95 Cretaceous Research pdfs, and I figure this is one of those historical volumes it's good to have, like Sibley and Ahlquist's bird phylogeny book.<br />
<br />
The format is an encyclopedia-style list of clades in phylogenetic order with Registration Number, Definition, Etymology, Reference Phylogeny, Composition, Apomorphies, Synonyms, Comments and Literature Cited. Rather like my Theropod Database, so no complaints there. Well, one complaint is really more to do with the PhyloCode itself where they decided to abbreviate definitions with the non-standard del/nabla triangle symbol ∇. If you want people to start using your format, you might want to choose symbols that exist on a standard keyboard. Alt+2207 is supposed to generate it in Windows, but results in ƒ here in Blogger. Anyone know the correct Unicode numbers?<br />
<br />
On to the substance, where Phylonyms covers all life. Dinosaurs are the last section of the book, and non-avian dinosaurs get all of four definitions-<br />
<br />
<br />
<br />
<b><i>Dinosauria </i></b>R. Owen 1842 [M. C. Langer, F. E. Novas, J. S. Bittencourt, M. D. Ezcurra, and J. A. Gauthier], converted clade name<br /><br /><b>Registration Number</b>: 194<br /><br /><b>Definition</b>: The smallest clade containing <i>Iguanodon bernissartensis</i> Boulenger in Beneden 1881 (Ornithischia/Euornithopoda) <i>Megalosaurus bucklandii</i> Mantell 1827 (Theropoda/Megalosauroidea) and <i>Cetiosaurus oxoniensis</i> Phillips 1871 (Sauropodomorpha).<br />
<br />
I'm glad we've standardized which theropod, ornithischian and sauropodomorph are used (or so I thought, see below), but otherwise there's not much to say. The caveats around which apomorphies are also found in <i>Nyasasaurus </i>and at least some silesaurs illustrate why apomorphy-based definitions are bad. The reference phylogeny for this and Saurischia is Lloyd et al.'s (2008) supertree, which is quite outdated and has a lot of artifacts from being a supertree.<br />
<br />
<br />
<br />
<i><b>Saurischia</b></i> H. G. Seeley 1888 [J. A. Gauthier, M. C. Langer, F. E. Novas, J. Bittencourt, and M. D. Ezcurra], converted clade name<br /><br /><b>Registration Number</b>: 195<br /><br /><b>Definition</b>: The largest clade containing <i>Allosaurus fragilis</i> Marsh 1877 (Theropoda/Carnosauria) and <i>Camarasaurus supremus</i> Cope 1877 (Sauropodomorpha), but not <i>Stegosaurus stenops</i> Marsh 1887 (Ornithischia/Stegosauridae).<br />
<br />
It's rather odd the same authors didn't choose the same specifiers for each dinosaurian clade as they did in the previous definition, leaving us without a neat node-stem triplet. Instead they went with the Kischlatian approach of using taxa"mentioned and figured as examples of their respective groups by Seeley (1888)." This is funny because I don't think this rationale is ever suggested in the PhyloCode, whereas Dinosauria and Saurischia are actually the official examples used for Recommendation 11F encouraging node-stem triplets ("If it is important to establish two names
as applying to sister clades regardless of the phylogeny, reciprocal
maximum-clade definitions should be used in which the single internal
specifier of one is the single external specifier of the other, and vice
versa"). Specifically- "If one wishes to define the names <em class="nomen">Saurischia</em> and <em class="nomen">Ornithischia</em> such that they will always refer to sister clades, <em class="nomen">Saurischia</em> might be defined as the largest clade containing <em class="nomen">Megalosaurus bucklandii</em> <span class="author">Mantell</span> 1827 but not <em class="nomen">Iguanodon bernissartensis</em> <span class="author">Boulenger</span> in <span class="author">Beneden</span> 1881, and <em class="nomen">Ornithischia</em> would be defined as the largest clade containing <em class="nomen">Iguanodon bernissartensis</em> but not <em class="nomen">Megalosaurus bucklandii</em>. To stabilize the name <em class="nomen">Dinosauria</em> as referring to the clade comprising <em class="nomen">Saurischia</em> and <em class="nomen">Ornithischia</em>, <em class="nomen">Dinosauria</em> should be defined as the smallest clade containing <em class="nomen">Megalosaurus bucklandii</em> and <em class="nomen">Iguanodon bernissartensis</em>." <br />
<br />Ornithoscelida and its consequences are mentioned, but I'm glad more time is not taken up with it as I expect the hypothesis to fall away as Baron's phylogenetic mistakes are not followed by future authors.<br />
<br />
<br />
<br />
<b><i>Sauropodomorpha </i></b>F. R. von Huene 1932 [M. Fabbri, E. Tschopp, B. McPhee, S. Nesbitt, D. Pol, and M. Langer], converted clade name<br /><br /><b>Registration Numbe</b>r: 295<br /><br /><b>Definition</b>: The largest clade containing <i>Saltasaurus loricatus</i> Bonaparte and Powell 1980 (Sauropodomorpha) but not <i>Allosaurus fragilis</i> Marsh 1877 (Theropoda) and <i>Iguanodon bernissartensis</i> Boulenger in Beneden 1881 (Ornithischia).<br />
<br />
I dislike the use of <i>Saltasaurus </i>as the internal specifier, which is a holdover of Sereno's weird use of deeply nested OTUs when others would be more historically relevant and/or eponymous. Fabbri et al. defend the choice because "Fossil specimens referred to <i>Saltasaurus loricatus</i> are abundant, the species is well known, and its phylogenetic position is consistent among phylogenetic analyses", but this would be even more true for e.g. <i>Camarasaurus supremus</i> used in Saurischia's definition. The other specifiers are a mix of those in Dinosauria's and Saurischia's definition, so there's absolutely no consistency. The reference phylogeny is Otero et al.'s (2015) <i>Sefapanosaurus </i>description using Yates' matrix, so is fine.<br /><br />There's a rare error in the comments for this entry. Fabbri et al. state "<i>Segnosaurus galbinensis</i> from the Cretaceous was briefly thought to be a relatively early diverging sauropodomorph (Paul, 1984; Gauthier, 1986; Olshevsky, 1991). More material referable to that species and the discovery of closely related taxa later showed that <i>Segnosaurus galbinensis</i> is part of the Therizinosauria", but material of <i>S. galbinensis</i> besides that initially recovered in the 1970s is not known.<br />
<br />
<br />
<br />
<i><b>Theropoda </b></i>O. C. Marsh 1881 [D. Naish, A. Cau, T. R. Holtz, Jr., M. Fabbri, and J. A. Gauthier], converted clade name<br /><br /><b>Registration Number</b>: 216<br /><br /><b>Definition</b>: The largest clade containing <i>Allosaurus fragilis</i> Marsh 1877 (Theropoda) but neither <i>Plateosaurus engelhardti</i> Meyer 1837 (Sauropodomorpha) nor <i>Heterodontosaurus tucki</i> Crompton and Charig 1962 (Ornithischia).<br />
<br />
Here we've chosen two completely different specifiers for Sauropodomorpha and Ornithischia, so again we have no consistency. The reference phylogeny is Cau (2018), which is ideal. <br />
<br />
<br />
<br />
What about the rest? It's a HUGE volume, and obviously most of Pan-Biota is outside my area of expertise. One obvious issue is the wildly varying coverage of different clades. Apparently nobody could be bothered with the vast majority of vertebrates (euteleosts) or animals (insects, except one definition for Trichoptera), and Molluska doesn't even get a definition. But we do get several entries for edrioasterid taxa down to subfamily-level, generally obscure Paleozoic echinoderms. Closer to dinosaurs, there's nothing at all for pan-crocs, but we get an entry for Pterosauromorpha for which only <i>Scleromochlus </i>is given as a plausible non-pterosaurian example (perhaps wrongly- Bennett, 2020). <br />
<br />
Then there are the apomorphy-based definitions which will cause headaches in the future. Look at Apo-Chiroptera- "Definition: The clade for which the unique modifications of the hand, forearm, humerus, scapula, hip, and ankle (see Diagnostic Apomorphies) associated with flapping flight, as inherited by <i>Vespertilio murinus</i> Linnaeus 1758, are apomorphies." Then you go down to the nine listed sets of Diagnostic Apomorphies like "Modification of the scapula: Scapular spine originates at the posterior edge of the glenoid fossa. Long axis of scapular spine offset 20–30 degrees from axis of rotation of the humeral head. Scapular spine reduced in height—acromion process appears more strongly arched and less well supported than in other mammals. Presence of at least two facets in infraspinous fossa." These are all going spread out as more stem bats are discovered, and indeed the authors already note "Simmons and Geisler (1998) included the absence of claws on wing digits III-V with this suite of modifications; however, the presence of claws on all the wing digits of <i>Onychonycteris </i>suggests that claws were present primitively in Apo-Chiroptera."<br />
<br />
Ungulata is defined by Archibald as "The least inclusive crown clade containing <i>Bos primigenius</i> Bojanus 1827 (= <i>Bos taurus</i> Linnaeus 1758) (Artiodactyla) and <i>Equus ferus</i> Boddaert 1785 (= <i>Equus caballus</i> Linnaeus 1758) (Perissodactyla), provided that this clade does not include <i>Felis silvestris</i> Schreber 1777 (= <i>Felis catus</i> Linnaeus 1758) (Carnivora), <i>Manis pentadactyla</i> Linnaeus 1758 (Pholidota), <i>Vespertilio murinus</i> Linnaeus 1758 (Chiroptera), or <i>Erinaceus europaeus</i> Linnaeus 1758 (Lipotyphla)." But this doesn't exist in molecular studies, including those of ultraconserved elements, which consistently place carnivorans, pangolins and bats closer to perrisodactyls. So this is likely to be a historical footnote, as well established molecular relationships end up trumping morphological relationships in every example I know of.<br />
<br />
Finally, we get Pan-Lepidosauria for the total group of lepidosaurs, which has been Lepidosauromorpha for over thirty years. Yet Archosauromorpha is retained as "The least inclusive clade containing <i>Gallus </i>(originally <i>Phasianus</i>) <i>gallus </i>(Aves) (Linnaeus 1758), <i>Alligator </i>(originally <i>Crocdilus</i>) <i>mississippiensis </i>(Daudin 1802) (Crocodylia), <i>Mesosuchus browni</i> Watson 1912 (Rhynchosauria), <i>Trilophosaurus buettneri</i> Case 1928 (Trilophosauridae), <i>Prolacerta broomi</i> Parrington 1935 (Prolacertiformes), and <i>Protorosaurus speneri</i> von Meyer 1830 (Protorosauria)" even though Pan-Archosauria is also used for the total group of archosaurs, traditionally the definition of Archosauromorpha. I agree our new Archosauromorpha deserved a name for being a generally recognized group, whereas whether e.g. choristoderes or sauropterygians fell out closer to lizards or birds is highly unstable. But I would have rather kept the tradition of -omorpha for the stem clades and gave this a new name.<br />
<br />
<br />
<br />
Overall, I'm not very impressed for something 20 years in the making that intends to be so important. How do you contradict your own example for choosing specifiers in four papers, where two share the same author list, the other two share another author (Fabbri), and each of those shares an author with both of the first two (Langer and Gauthier)? And one of those is an editor for the volume. Nothing could be negotiated in over two decades? But it's what we have to work with now, and in the name of consistancy I'll adopt the definitions proposed. Now to see what happens when RegNum goes online.<br />
<br />
<b>References</b>- Lloyd, Davis, Pisani, Tarver, Ruta, Sakamoto, Hone, Jennings and Benton, 2008. Dinosaurs and the Cretaceous terrestrial revolution. Proceedings of the Royal Society B. 275, 2483-2490.<br /><br />Otero, Krupandan, Pol, Chinsamy and Choiniere, 2015. A new basal sauropodiform from South Africa and the phylogenetic relationships of basal sauropodomorphs. Zoological Journal of the Linnean Society. 174, 589-634.<br />
<br />
<span style="color: black;"><span style="font-family: inherit, Times New Roman, serif;">Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Società Paleontologica Italiana. 57(1),
1-25.</span></span><br />
<br />
<span style="color: black;"><span style="font-family: inherit, Times New Roman, serif;">Bennett, 2020. Reassessment of the Triassic archosauriform <i>Scleromochlus taylori</i>: Neither runner nor biped, but hopper. PeerJ. 8:e8418.</span></span><br />
<br />
de Queiroz, Cantiono and Gauthier, 2020. <span>Phylonyms: A Companion to the PhyloCode, 1st Edition. Taylor & Francis Group. 1323 pp.</span> <br />
<br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com7tag:blogger.com,1999:blog-3248412803814730250.post-11597394981903525252020-03-16T04:31:00.000-07:002020-03-16T04:40:13.389-07:00What is Oculudentavis if it's not a theropod?In my last post, I argued the recently described <i>Oculudentavis </i>(Xing et al., 2020) is not a theropod. So what is it? To answer that question, I entered it into Simoes et al.'s (2018) sauropsid analysis which emphasizes basal lepidosauromorphs and comes out with basal gekkos and nested iguanians even using just morphological characters. To test Jingmai's avialan hypothesis, I also added <i>Archaeopteryx </i>to the matrix. The result is 384 MPTs of 2337 steps each.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiYDHNLsgLcsmdnJX0V5ORac5Vp4GBAG_TPH7gkI3HL9RM3TPaDtTu9Ayi60q8NJXCgGjc1A-XTydZQwCF7gw_n30_9Ss_MjTgea2OkUZqbZFv0R4ttR2uwYlG3sPCZbM0qgIejpHJzjKI/s1600/Oculudentavis+phylo.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1411" data-original-width="1107" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiYDHNLsgLcsmdnJX0V5ORac5Vp4GBAG_TPH7gkI3HL9RM3TPaDtTu9Ayi60q8NJXCgGjc1A-XTydZQwCF7gw_n30_9Ss_MjTgea2OkUZqbZFv0R4ttR2uwYlG3sPCZbM0qgIejpHJzjKI/s320/Oculudentavis+phylo.jpg" width="251" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Strict consensus of 384 MPTs of Simoes et al.'s (2018) analysis after adding <i>Oculudentavis </i>and <i>Archaeopteryx</i>. Compare to Extended Data Figure 3 of Simoes et al..</td></tr>
</tbody></table>
<div class="separator" style="clear: both; text-align: center;">
</div>
As you can see, <i>Oculudentavis </i>resolves as a stem-squamate in a trichotomy with <i>Huehuecuetzpalli</i> and squamates, while <i>Archaeopteryx </i>is an archosauromorph sister to <i>Erythrosuchus</i>. And this matrix didn't score for scleral ossicle shape, posttemporal fenestra size or maxillary tooth row length. After scoring <i>Oculudentavis</i>, its teeth are clearly not acrodont, it seems to have a ventral parietal fossa and lacks an ossified laterosphenoid. The authors could have made it easier to evaluate by separating the cranial elements in the 3D pdf file. As it is, a lot of palatal and braincase info is uncertain. But <i>Huehuecuetzpalli </i>is Albian compared to <i>Oculudentavis</i>' Cenomanian, and has a skull length of 32 mm (19 mm in the juvenile) versus 14 mm in <i>Oculudentavis</i>.<br />
<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVoLatRWzCRb9TVLhlTPFVLIzVP__AcphoUpuTpF7bF8VnExSomzhi7-G62FVDjKrQAZJz5YpR04AwvKd0qEbxwzLq4KadOyqtt1At-Eu0KDBBv4MpS_f7kBkPszRS37lZobJu0BlvhYI/s1600/Oculudentavis+comp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1053" data-original-width="749" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVoLatRWzCRb9TVLhlTPFVLIzVP__AcphoUpuTpF7bF8VnExSomzhi7-G62FVDjKrQAZJz5YpR04AwvKd0qEbxwzLq4KadOyqtt1At-Eu0KDBBv4MpS_f7kBkPszRS37lZobJu0BlvhYI/s320/Oculudentavis+comp.jpg" width="227" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Huehuecuetzpalli </i>skull (top; after Reynoso, 1998), <i>Oculudentavis </i>skull and separate mandible (middle; after Xing et al., 2020), and <i>Archaeopteryx </i>skull (after Rauhut, 2014).</td></tr>
</tbody></table>
<br />
<b>References</b>- <span style="color: #222222;">Reynoso, 1998. <i>Huehuecuetzpalli mixtecus</i> gen. et sp. nov: A basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodriguez, central Mexico. </span><span style="color: #222222;"><span class="acknowledgment-journal-title">Philosophical Transactions of the Royal Society B: Biological Sciences. 353, 477-500.</span></span><br />
<span style="color: #222222;"><span class="acknowledgment-journal-title">Rauhut, 2014. New observations on the skull of <i>Archaeopteryx</i>. Paläontologische
Zeitschrift. 88(2), 211-221.<br /><br />Simōes, Caldwell, Talanda, Bernardi, Palci, Vernygora, Bernardini, Mancini and Nydam, 2018. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature. 557(7707), 706-709.</span></span><span class="reference-text"><cite class="citation journal"></cite></span><br />
<span style="color: #222222;"><span class="acknowledgment-journal-title"></span></span><br />
<span style="color: #222222;"><br /></span>
<span style="color: #222222;">Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li,<i>
</i>2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature.
579, 245-249.</span>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com9tag:blogger.com,1999:blog-3248412803814730250.post-64491945007810244422020-03-12T04:02:00.001-07:002020-03-12T04:02:32.404-07:00Oculudentavis is not a theropodHi all. This week we got the announcement of a tiny theropod skull in Myanmar amber, which was bound to happen eventually as amazing finds from that deposit keep being published. Alas, whatever <i>Oculudentavis </i>is, it's not a theropod.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJVC9ylsdQcR_2w-8GDn8CdnxleUXrGgCnOQEh9Mt4YChyphenhyphenQza474GBW83QQXgzAXf_dF489DLt9o1HjtyoHU6oyk01R8oTxlYfx59pfIivwof4kMKpq1CZivtfOfssvhaZu4EY1KIHMZk/s1600/Oculudentavis+main.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1037" data-original-width="1416" height="234" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJVC9ylsdQcR_2w-8GDn8CdnxleUXrGgCnOQEh9Mt4YChyphenhyphenQza474GBW83QQXgzAXf_dF489DLt9o1HjtyoHU6oyk01R8oTxlYfx59pfIivwof4kMKpq1CZivtfOfssvhaZu4EY1KIHMZk/s320/Oculudentavis+main.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Oculudentavis </i>skull (after Xing et al., 2020).</td></tr>
</tbody></table>
<br />
Just look at it. No antorbital fenestra, incomplete ventral bar to the laterotemporal fenestra, huge posttemporal fenestrae, teeth that extend posteriorly far under the orbit...<br />
<br />
All of which might be coincidental, but then look at the mandible.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiLdxsEMPr04QLgNLl5vEAM8ZotL2Pkq7tYP3GmmL8gG7RiI3nde6KW-7peaNE9H5VJQlc3jbhNX1lvNcx3R3uo26IBAukr_XivQ4vByecmkca_oSfhyphenhyphennutKfdtHfBnReJeKvLNf_-TCLw/s1600/Oculudentavis+mandible.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="679" data-original-width="526" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiLdxsEMPr04QLgNLl5vEAM8ZotL2Pkq7tYP3GmmL8gG7RiI3nde6KW-7peaNE9H5VJQlc3jbhNX1lvNcx3R3uo26IBAukr_XivQ4vByecmkca_oSfhyphenhyphennutKfdtHfBnReJeKvLNf_-TCLw/s320/Oculudentavis+mandible.jpg" width="247" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Oculudentavis </i>mandible (after Xing et al., 2020).</td></tr>
</tbody></table>
<br />
That spike-like coronoid process is classic lepidosaur, plus the dentary is way too long compared to the post-dentary elements, then the description says "The tooth geometry appears to be acrodont to pleurodont; no grooves or sockets are discernable." And of course "the scleral ring is very large and is formed by elongated spoon-shaped ossicles; a morphology similar to this is otherwise known only in lizards (for example, <i>Lacerta viridis</i>)."<br />
<br />
Add to this the size of this partially fused specimen being smaller than any extant bird (14 mm), and no feather remains, and why is this a theropod again? The endocast is big, but why not a clade of brainier lizards or late surviving megalancosaurs by the Cenomanian?<br />
<br />
The authors add it to Jingmai's bird analysis where it ends in a huge polytomy closer to Aves than <i>Archaeopteryx</i>, but outside <a href="https://theropoddatabase.blogspot.com/2016/01/how-not-to-redefine-ornithuromorpha.html" target="_blank">fake Ornithuromorpha</a>. That's often what happens when a taxon is wrongly placed in a clade. Note the figured placement between <i>Archaeopteryx </i>and <i>Jeholornis </i>is only found using implied weights. At least add it to e.g. Nesbitt's or Ezcurra's archosauromorph analyses, or Cau's theropod analyses before assuming it's a bird.<br />
<br />
Thanks to Ruben Molina Perez for suggesting this issue in the first place. <br />
<br />
<b>Reference</b>- <span style="color: #222222;">Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi and Li,<i>
</i>2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature.
579, 245-249.</span>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com15tag:blogger.com,1999:blog-3248412803814730250.post-72429136568585007402020-01-20T02:05:00.000-08:002020-01-27T13:10:56.417-08:00Details on Teinurosaurus and random musingsHi all. When updating The Theropod Database I noticed my entry for <i>Teinurosaurus </i>is pathetically bad- wrong authors, wrong age, wrong size, and generally missing the complicated history of this innocuous vertebra. How embarrassing! So here's the revised version that will be uploaded-<br />
<br />
<b><i>Teinurosaurus</i></b> Nopcsa, 1928<br />
= <span style="font-style: italic;">Saurornithoides</span> Nopcsa, 1928 (preoccupied Osborn, 1924)<br />
= <i>Caudocoelus</i> Huene, 1932<br />
<i><b>T. sauvagei</b></i> (Huene, 1932) Olshevsky, 1978<br />
= <i>Caudocoelus sauvagei</i> Huene, 1932<br />
<b>Tithonian, Late Jurassic<br />Mont-Lambert Formation, Hauts-de-France, France</b><br />
<b>Holotype</b>- (BHN2R 240; = Boulogne Museum 500) incomplete distal caudal vertebra (75 mm)<br />
<span style="font-weight: bold;">Diagnosis</span>- Provisionally indeterminate relative to <span style="font-style: italic;">Kaijiangosaurus</span>, <span style="font-style: italic;">Tanycolagreus</span> and <span style="font-style: italic;">Ornitholestes</span>.<br />
<span style="font-weight: bold;">Other diagnoses</span>- (after Huene, 1932; compared to <span style="font-style: italic;">Elaphrosaurus</span>) centrum wider; narrower ventral surface; ventral median groove wider; transversely narrower prezygapophyses.<br />
While Huene attmpted to distinguish <span style="font-style: italic;">Teinurosaurus</span> from <span style="font-style: italic;">Elaphrosaurus</span>,
only the wider median ventral groove is apparent in existing photos of
the former. This is compared to the one distal caudal of the
latter figured in ventral view, but as Kobayashi reports grooves become
distally narrower in <span style="font-style: italic;">Harpymimus</span> while Ostrom reports they become distally wider in <span style="font-style: italic;">Deinonychus</span>,
groove width is not considered taxonomically distinctive at our current
level of understanding. Indeed, this lack of data is most
relevent to both diagnosing and identifying <span style="font-style: italic;">Teinurosaurus</span>.
Very few taxa have detailed descriptions of distal caudal vertebrae or
more than lateral views figured, let alone indications of variation
within the distal caudal series. So the facts that <span style="font-style: italic;">Fukuiraptor</span> and <span style="font-style: italic;">Deinonychus</span> share ventrally concave central articulations with <span style="font-style: italic;">Teinurosaurus</span> in their single anteriorly/posteriorly figured distal caudal vertebra, or that <span style="font-style: italic;">Afromimus</span>, "Grusimimus" and <span style="font-style: italic;">Falcarius</span>
also have have wide ventral grooves in their few ventrally figured
distal caudals, are not considered taxonomically important. <br />
<span style="font-weight: bold;">Comments</span>- Sauvage (1897-1898;
in a section written in January 1898) first mentioned a distal caudal
vertebra he referred to the ornithischian <span style="font-style: italic;">Iguanodon prestwichii</span> (now recognized as the basal styracosternan <span style="font-style: italic;">Cumnoria prestwichii</span>) - "<span class="tlid-translation translation" lang="en"><span class="" title="">We
are disposed to regard as belonging to the same species the caudal
vertebra of a remote region, the part which we figure under n ° 7, 8"
[translated].</span></span> Note Galton (1982) was incorrect in
claiming Sauvage reported on this specimen in his 1897 paper (written
December 6), which includes a section on <span style="font-style: italic;">prestwichii</span>
nearly identical to the 1897-1898 one but which lacks the paragraph
describing this vertebra. This could provide a specific date of
December 1897 to January 1898 for the discovery and/or recognition of
the specimen. Huene (1932) correctly noted Sauvage mislabeled
plate VII figure 8 as dorsal view, when it is in ventral view as
understood by the text. Compared to <span style="font-style: italic;">Cumnoria</span>,
the caudal is more elongate (length 3.93 times posterior height
compared to 2.54 times at most), has a ventral median groove instead of
a keel, and the prezygapophyseal base in 71% of the anterior central
height compared to ~30-40%, all typical of avepods. Nopcsa (1928)
recognized its theropod nature and in his list of reptile genera meant
to use a footnote to propose <span style="font-style: italic;">Teinurosaurus</span>
as a "new name for the piece described and figured by Sauvage (Direct.
Traveaux Geol. Portugal Lisbonne 1897-1898, plate VII, Fig. 7-10) as
late caudal of <span style="font-style: italic;">Iguanodon Prestwichi</span>." <span style="font-style: italic;">Teinurosaurus</span>
is listed as an aublysodontine megalosaurid (not as an ornithomimine,
contra Galton), roughly equivalent to modern Eutyrannosauria.
However due to a typographical error, the footnote's superscript 1 was
placed after <span style="font-style: italic;">Saurornithoides</span> instead of <span style="font-style: italic;">Teinurosaurus</span>. Sauvage (1929) corrected this in an addendum- "footnote 1 does not refer to <span style="font-style: italic;">Saurornithoides</span> (line 19 from below) but to <span style="font-style: italic;">Teinurosaurus</span>
(last line of text)." Unfortunately, Huene missed the addendum,
and thus wrote "Nopcsa recognized in 1927 (43, p. 183) that this was a
coelurosaur and intended to give it a name, but used one already used
by Osborn, namely "<span style="font-style: italic;">Saurornithoides</span>" (91, 1924, p. 3- 7). For this reason, a new name had to be given here" [translated]. Huene's proposed new name was <span style="font-style: italic;">Caudocoelus sauvagei</span>, placed in Coeluridae and "somewhat reminiscent of <span style="font-style: italic;">Elaphrosaurus</span>."
Huene is also perhaps the first of several authors to place the
specimen in the Kimmeridgian, when it is actually from the Tithonian
(Buffetaut and Martin, 1993; as Portlandian). Galton wrote
"Lapparent and Lavocat (1955: 801) gave a line drawing of the vertebra
after Sauavage (1898) and included it in the section on <span style="font-style: italic;">Elaphrosaurus</span>" and that the specimen "was referred to <span style="font-style: italic;">Elaphrosaurus</span>
by Lapparent and Lavocat (1955)." This was perhaps done because
Huene explicitly compared the two, ironically making it the only taxon
distinguished from <span style="font-style: italic;">Teinurosaurus</span> at the time. Most of Huene's characters cannot be checked in the few published photos of <span style="font-style: italic;">Teinurosaurus</span>, but the ventral median sulcus is indeed much wider than <span style="font-style: italic;">Elaphrosaurus</span>. Ostrom (1969) was the first author to detail Nopcsa's (1929) addendum, stating "Nopcsa's name <span style="font-style: italic;">Teinurosaurus</span> has clear piority over Huene's <span style="font-style: italic;">Caudocoelus</span>, but since Nopcsa failed to provbide a specific name, <span style="font-style: italic;">Teinurosaurus</span> is not valid." Olshevsky (1978) solved this by writing "<span style="font-style: italic;">Teinurosaurus</span> has clear priority over <span style="font-style: italic;">Caudocoelus</span>, as noted in Ostrom 1969, and it is certainly a valid generic name. The species <span style="font-style: italic;">Caudocoelus sauvage</span>i is proposed here as the type species of the genus <span style="font-style: italic;">Teinurosaurus</span>, resulting in the new combination <span style="font-style: italic;">Teinurosaurus sauvagei</span>
(von Huene 1932) as the proper name of the type specimen." He
also claimed "the specimen itself, unfortunately, was destroyed during
World War II and thus must remain a nomen dubium." This was
repeated by Galton, but as Buffetaut et al. (1991) wrote- "Contrary to
a widespread opinion (expressed, for instance, by Lapparent, 1967), the
vertebra in question has survived two world wars and years of neglect,
like a large part of the other fossil reptile remains in the
collections of the Boulogne Natural History Museum (see Vadet and Rose,
1986)." Olshevsky noted Steel misunderstood Nopsca in a different
way, believing <span style="font-style: italic;">Teinurosaurus</span> instead of <span style="font-style: italic;">Aublysodon</span> was a "name, proposed by Cope in 1869 ... used instead of <span style="font-style: italic;">Deinodon</span>", as stated under superscript 2. Galton did have the first modern opinion on <span style="font-style: italic;">Teinurosaurus</span>' affinities, stating "In addition to <span style="font-style: italic;">Elaphrosaurus</span>, elongate prezygapophyses occur in the allosaurid <span style="font-style: italic;">Allosaurus</span> and the dromaeosaurid <span style="font-style: italic;">Deinonychus</span>,
so this caudal vertebra can only be identified as theropod, family
incertae sedis." Buffetaut and Martin (1993) agreed, saying "no
really distinctive characters that would allow a familial assignment
can be observed." Ford (2005 online) gave the type repository as
"Dortigen Museum", but this is a misunderstanding based on Huene's
"Boulogne-sur-mer (Nr. 500 im dortigen Museum)", which
translated is "Boulogne-sur-mer (No. 500 in the museum there)",
referring to the Boulogne Museum where it has always been held.
It was originally number 500, but was recatalogued at some point.<br />
Sauvage lists the vertebra's length as 75 mm and his plate at natural
size would have it be 79 mm, Huene lists it as 11 cm (110 mm) and his
figure at 1:2 size would have it be 152 mm. Galton's drawing with
supposed 5 cm scale would have it be 235 mm, while Buffetaut and
Martin's plate with scale would leave it at 74 mm. As Huene's and
Galton's figures are taken from Sauvage's original plate and the newest
and unique photo matches Sauvage's reported length almost exactly, 75
mm is taken as the correct length.<br />
<span style="font-weight: bold;">Relationships</span>- While prior authors haven't specified <span style="font-style: italic;">Teinurosaurus</span>' relationships past Theropoda (besides Lapparent and Lavocat's apparent synonymy with <span style="font-style: italic;">Elaphrosaurus</span>),
there are several ways to narrow down its identity. Only
neotheropods are known from the Late Jurassic onward, so
coelophysoid-grade taxa are excluded. Some theropod clades were
too small to have a 75 mm caudal, including most non-tyrannosauroid
coelurosaurs besides ornithomimosaurs, therizinosaurs and
eudromaeosaurs. The former two are unknown from the Jurassic, and
additionally paravians like eudromaeosaurs lack any neural spine by the
time the centrum gets as elongate as <span style="font-style: italic;">Teinurosaurus</span> (e.g. by caudal 12 in <span style="font-style: italic;">Deinonychus</span> at elongation index of 2.4). <span style="font-style: italic;">Teinurosaurus</span> has an elongation index (centrum length/height) of 3.9, which also excludes Ceratosauridae, <span style="font-style: italic;">Beipiaosaurus</span>
+ therizinosauroids and oviraptorosaurs. Prezygapophyses basal
depth is significantly less in ceratosaurids, megalosaurids, carnosaurs
except <span style="font-style: italic;">Neovenator</span>, compsognathids, <span style="font-style: italic;">Fukuivenator</span> and <span style="font-style: italic;">Falcarius</span>. Remaining taxa are elaphrosaur-grade ceratosaurs, piatnitzkysaurids, <span style="font-style: italic;">Neovenator</span> and basal tyrannosauroids. <br />
<b>References</b>- Sauvage, 1897. Notes sur les Reptiles Fossiles (1). Bulletin de la Société géologique de France. 3(25), 864-875.<br />
Sauvage, 1897-1898. Vertebres Fossiles du Portugual, Contributions
a l'etude des poissions et des reptiles du Jurassique et du Cretaceous. Direction
des Travaux Geologiques Portugal. 1-46.<br />
Osborn, 1924. Three new Theropoda, <i>Protoceratops</i> zone, central Mongolia.
American Museum Novitates. 144, 1-12.<br />
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1,
163-188.<br />
<span class="reference-text">Nopcsa, 1929. Addendum "The genera of reptiles". </span>Palaeobiologica. 2, 201.<br />
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), 361 pp. <br />
Lapparent and Lavocat, 1955. Dinosauriens. In Piveteau (ed.). Traite de Paleontologie. Masson et Cie. 5, 785-962.<br />
Lapparent, 1967. Les dinosaures de France. Sciences. 51, 4-19.<br />
Ostrom, 1969. Osteology of <i>Deinonychus antirrhopus</i>, an unusual theropod
from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin.
30, 1-165. <br />
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag. 87 pp. <br />
Olshevsky, 1978. The archosaurian taxa (excluding the Crocodylia). Mesozoic
Meanderings. 1, 50 pp.<br />
Galton, 1982. <i>Elaphrosaurus</i>, an ornithomimid dinosaur from the Upper
Jurassic of North America and Africa. Paläontologische Zeitschrift. 56, 265-275.<br />
Vadet and Rose, 1986. Catalogue commente des types et figures de dinosauriens, ichthyosauriens, sauropterygiens, pterosauriens et cheloninens du Musée d'Histoire Naturelle de Boulogne-sur-Mer. In E. Buffetaut, Rose and Vadet (eds.). Vértébrés Fossiles du Boulonnais. Mémoires de la Société Académique du Boulonnais. 1(2), 85-97.<br />
Rose, 1987. Redecouverte d'une vertebre caudale reptilienne (Archosauriens) de status controverse et provenant des terrains jurassiques superieurs du Boulonnais. Bulletin de la Société académique du Boulonnais. 1(5), 150-153.<br />
Buffetaut, Cuny and le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1-2), 17-42.<br />
Buffetaut and Martin, 1993. Late Jurassic dinosaurs from the Boulonnais (northern France): A review. Revue de Paléobiologie. 7(vol. spéc.),
17-28.<br />
Ford, 2005 online. <a href="http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp">http://www.paleofile.com/Dinosaurs/Theropods/Teinurosaurus.asp</a><br />
<br />
<b>* minor edits (see Marjanovic's comment)</b><br />
<br />
And before we go, here are a couple more tidbits I've noticed in the upcoming update...<br />
<br />
- <span class="tlid-translation translation" lang="en">That theropod tail preserved in Burmese amber (DIP-V-15103) described by Xing et al. (2016) was only placed as specifically as a non-pygostylian maniraptoriform. But as the deposits are Gondwanan (e.g. Poinar, 2018), the range of potential Cenomanian theropods is better understood. And only one group has caudal centra over three times longer than tall- unenlagiines. I bet DIP-V-15103 is our first sample of preserved plumage in an unenlagiine, which makes you wonder if the weird alternating barb placement was a feature that evolved on Gondwana, and if so did <i>Rahonavis</i>' remiges exhibit it too?</span><br />
<span class="tlid-translation translation" lang="en"><br /></span>
<span class="tlid-translation translation" lang="en">- Does anyone realize both "Tralkasaurus" (Cerroni et al., 2019) and "Thanos" (Delcourt and Iori, 2018) are nomina nuda? Neither are in an official volume yet, though "Tralkasaurus" is scheduled for March and "Thanos" will probably make it this year if the average papers per volume of Historical Biology holds up. "Tralkasaurus" has an empty space in its "Zoobank registration:" section, while the "Thanos" paper doesn't mention ZooBank at all, and neither </span><span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en">show up in ZooBank searches. Also, one of "Thanos"' supposed autapomorphies is a deep </span></span><span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en">prezygapophyseal spinodiapophyseal fossa, which does not exist in abelisaurs as it would require a spinodiapophyseal lamina. The labeled structure seems internal, </span></span><span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en">probably the centroprezygapophyseal fossa or prezygapophyseal
centrodiapophyseal fossa based on CT-scanned noasaurid cervical
DGM929-R. That leaves axial pleurocoel size and distance from each other, and ventral keel strength as suggested characters. Which can only be compared to <i>Carnotaurus </i>among brachyrostrans. Hmmm...</span></span><br />
<span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en"><br /></span></span>
<span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en"><b>References</b>- </span></span><span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en">Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang,
Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur tail with
primitive plumage trapped in Mid-Cretaceous amber. Current Biology.
26(24), 3352-3360.</span></span><br />
<span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en"><br />Delcourt and Iori, 2018. A new Abelisauridae (Dinosauria: Theropoda)
from São José do Rio Preto Formation, Upper Cretaceous of Brazil and
comments on the Bauru Group fauna. Historical Biology. DOI:
10.1080/08912963.2018.1546700<br /><br />Poinar, 2018. Burmese amber: Evidence of Gondwanan origin and Cretaceous dispersion. Historical Biology. DOI: 10.1080/08912963.2018.1446531</span></span><br />
<br />
<span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en"></span></span>Cerroni, Motta, Agnolín, Aranciaga Rolando, Brissón Egli and Novas,
2019. A new abelisaurid from the Huincul Formation
(Cenomanian-Turonian; Upper Cretaceous) of Río Negro province,
Argentina. Journal of South American Earth Sciences. 98, 102445.<br />
<span class="tlid-translation translation" lang="en"><span class="tlid-translation translation" lang="en"></span></span>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com4tag:blogger.com,1999:blog-3248412803814730250.post-79036948430414855122020-01-02T08:50:00.000-08:002020-01-02T08:50:00.463-08:00Happy New Year 2020Hi all. A <a href="https://www.theropoddatabase.com/Updates.htm" target="_blank">Theropod Database update</a> is online, with the main additions being troodontid information and info from the Hayashibara Museum of Natural Sciences Research Bulletins 1-3. I love these publications and wish more like them existed for other collections. They detail the expeditions into Mongolia with exact discovery dates and field numbers for taxa like <i>Nomingia</i>, <i>Elsornis </i>and <i>Aepyornithomimus</i>, and tons of still undescribed specimens. It's amazing just how many ornithomimosaurs are known from the Bayanshiree Formation for instance, when only the <i>Garudimimus </i>holotype has been described. There are over twenty more including the sort-of-described "<i>Gallimimus</i>" "mongoliensis" specimen IGM 100/14. So often for new taxa, especially those from the Jehol biota, no information is provided in the description as to when the specimen was discovered. I get that many are found by non-professionals and given to museums, but at least say "the specimen was given to the museum on x-x-xx by someone who said it was excavated around year y." Next up, halszkaraptorine and dromaeosaurid updates...<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjV5RDu9Awq6vpzO2ZTpveB2AjA7VMkvH_-j2ogg0w5SWzB7vc6YgtIjEfpv38cZXGyFIyPGEpzcOMnsPXUUDy6iSkwhprqEeZJz8rBaP1V7RgP-dLFZMmvO5A2ksI3iZ1ub5wOHjXun1M/s1600/Gobivenator+referred.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="497" data-original-width="692" height="229" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjV5RDu9Awq6vpzO2ZTpveB2AjA7VMkvH_-j2ogg0w5SWzB7vc6YgtIjEfpv38cZXGyFIyPGEpzcOMnsPXUUDy6iSkwhprqEeZJz8rBaP1V7RgP-dLFZMmvO5A2ksI3iZ1ub5wOHjXun1M/s320/Gobivenator+referred.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">undescribed ?<i>Gobivenator </i>skull (HMNS coll.; field number 940801 TS-I WTB) (after Tsogtbaatar and Chinzorig, 2010).</td></tr>
</tbody></table>
<b>Reference</b>- Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian
Paleontological Center: 2002–2008. Hayashibara Museum of Natural
Sciences Research Bulletin. 3, 155-166. Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5tag:blogger.com,1999:blog-3248412803814730250.post-33066640169023988432019-08-22T11:57:00.000-07:002019-08-22T15:50:02.436-07:00Therizinosaurs in the Lori matrixNext up are therizinosaurs. These are one of the best analyzed clades because I incorporated all of Zanno's (2010) characters, which is by far the largest and most recent analysis of the group until the Lori paper was published. The topology is- <br />
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<div class="separator" style="clear: both; text-align: center;">
<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEinjnSmiEUlSh9L1vo0k8lnpyrDXCB-4uYPrFNsdqCGmNPxCXC5jsu9cyKcp2yXc3_bIMYFThvxTYPEz1j32nG-0pJijSOb1kUbY659UicMbtG3yVGqh5IEpgZh3t1UX9WV23C3TIiM2So/s1600/theriz.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="264" data-original-width="619" height="136" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEinjnSmiEUlSh9L1vo0k8lnpyrDXCB-4uYPrFNsdqCGmNPxCXC5jsu9cyKcp2yXc3_bIMYFThvxTYPEz1j32nG-0pJijSOb1kUbY659UicMbtG3yVGqh5IEpgZh3t1UX9WV23C3TIiM2So/s320/theriz.jpg" width="320" /></a></div>
<br />
<i>Falcarius </i>is the most basal taxon shown of course, but <i>Martharaptor </i>was pruned a posteriori and can fall out anywhere in Therizinosauria outside the <i>Alxasaurus </i>plus <i>Segnosaurus </i>clade. I tried including <i>Thecocoelurus</i>, but the Lori matrix is pretty terrible when it comes to scoring single vertebrae-<br />
<br />
Thecocoelurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????(01)0? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????2???? ?????????? ?????????? ?????????? ?????????? ???????0?? ?????????? ?????????? ?????????? ?????1???? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??0??????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????<br />
<br />
<i>Jianchangosaurus </i>fell out in the same place as its original description, with Cau's (2018) placement in Alvarezsauroidea taking 18 more steps so is very unlikely. Mine is the only published matrix besides Senter (2011) and its derivatives to use information from the second <i>Beipaiosaurus </i>specimen, and incorporated photos from Zanno and the new paper on the holotype skull elements too.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjKsTuGoDS1qXuZKgoHPwK4Whnq079uc2uQUlBs09oNfUtzBZgrjUVFkUsV1UohUKrYloJkoEJvgJdivSRFXVfSutkm0kKfVcYJMPXwTGg8m0e3txxkIBRQkL6pkByzmUhT4OIIybBg75E/s1600/Beipiaosaurus+inexpectus+V+11559+104.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjKsTuGoDS1qXuZKgoHPwK4Whnq079uc2uQUlBs09oNfUtzBZgrjUVFkUsV1UohUKrYloJkoEJvgJdivSRFXVfSutkm0kKfVcYJMPXwTGg8m0e3txxkIBRQkL6pkByzmUhT4OIIybBg75E/s320/Beipiaosaurus+inexpectus+V+11559+104.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Beipiaosaurus inexpectus</i> holotype (IVPP V11559) cervical vertebra in dorsal view (courtesy of Zanno).</td></tr>
</tbody></table>
Zanno also provided photos of <i>Alxasaurus </i>and <i>Enigmosaurus</i>, and its depressing how much of the former is lost. <i>Enigmosaurus </i>rather famously was shown by Zanno to not resemble Barsbold's original illustration that was the only reference picture known for over two decades. Placing <i>Enigmosaurus </i>closer to <i>Segnosaurus </i>than <i>Neimongosaurus </i>or <i>Erliansaurus </i>as in Zanno's tree takes 4 more steps. Forcing <i>Enigmosaurus </i>and <i>Erlikosaurus </i>to be sister taxa to simulate the synonymy mentioned by Barsbold (1983) takes 4 steps, so seems unlikely. The duo moves between <i>Nanshiungosaurus </i>and the <i>Segnosaurus </i>plus <i>Nothronychus </i>clade. We were the first analysis to include <i>"Chilantaisaurus" zheziangensis</i>, which emerged in a polytomy with <span style="font-style: italic;">Alxasaurus</span>, <span style="font-style: italic;">Enigmosaurus</span> and therizinosaurids.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsAvhrUfzFlh2YIhi51qMmbelsYWoQJhkfd85JoFWjajS8eji842wHW9jUz_Nnzyb2gFAFGWOh7CiWQwPrC8pbH2ZCvxq-oFCpNXd2DMWbmohvj8pNk-dWp5_gbF6GneroFwA5-yaNOtU/s1600/Alxasaurus+elesitaiensis+007.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsAvhrUfzFlh2YIhi51qMmbelsYWoQJhkfd85JoFWjajS8eji842wHW9jUz_Nnzyb2gFAFGWOh7CiWQwPrC8pbH2ZCvxq-oFCpNXd2DMWbmohvj8pNk-dWp5_gbF6GneroFwA5-yaNOtU/s320/Alxasaurus+elesitaiensis+007.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Alxasaurus elesitaiensis</i> holotype (IVPP V88402a) chevrons in right lateral view (natural order reversed) (courtesy of Zanno).</td></tr>
</tbody></table>
As was the case with <i>Archaeornithomimus? bissektensis</i>, we didn't include the possible chimaera of Bissekty Therizinosauria as an OTU, unlike Sues and Averianov (2015). But if you do want to experiment with it, here's the scorings. It emerges in a polytomy in the <i>Suzhousaurus </i>plus <i>Therizinosaurus </i>clade of therizinosaurids. Btw, <i>Archaeornithomimus? bissektensis</i> does fall out most parsimoniously sister to <i>A. asiaticus</i> when all Bissekty material is used.<br />
<br />
'Bissekty-Therizinosauroidea' ????1??0?? ????1????? ?????1???? ?1???????? ?????{01}1000 ?01??????? 00???????? 2??21?0??? ???????00{123} {12}00(01)2011?? ??0???0(01)00 ??11????{12}? ?{01}0?100??? ?????????? ?1{01}?0????? ??0?0???{012}0 000(01)?010?? ?????????? ?????????? ?????0{12}?00 1?0???0??0 ??0??{01}0??? 0?1??????? ????0?1?0? {01}0?1???{01}?? ??0??????? ?????????? ?????????0 ???001??00 ??00?01?1? 10???????? ?????????1 ???0?????? ???1?????? ??????0??1 0???1(12)???? ????0???1? ????0?011? ???0????0? ???0{12}0???? ????????0? 0?0????1?? ?????????? ??010101?1 ??0?0(01)???? ?????{01}010? ???100???? ????????11 1010?????? ??0??0???? ?????????? ?????????? ??????0??? ??00?????? ?00??????? ?????????? ???????0-- -??00??010 ?????????? ??????-??? ???-010??1 0100????00 10?1?00??? ???0?00??? ?????????? ????0????? ???000???? ?0???-???? ?????????? ?????001??<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqt2wd0Ko3E9xCiW64AfFVVe24N98QBUsDmATjNxAvjye_2DSUkI3cGxSYIHKMRdCx64k7t_cCzQoAOoYlmWUwuRQfgw_Oszo52ADUnIEmRw4EJ4UFS98CFaFvYslAl4kxt1-6VJkCIjE/s1600/Enigmosaurus+sacrum+pelvis+ventral.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqt2wd0Ko3E9xCiW64AfFVVe24N98QBUsDmATjNxAvjye_2DSUkI3cGxSYIHKMRdCx64k7t_cCzQoAOoYlmWUwuRQfgw_Oszo52ADUnIEmRw4EJ4UFS98CFaFvYslAl4kxt1-6VJkCIjE/s320/Enigmosaurus+sacrum+pelvis+ventral.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Enigmosaurus mongoliensis </i>holotype (IGM 100/84) synsacrum and ilium in ventral view (courtesy of Zanno).</td></tr>
</tbody></table>
Next is Therizinosauridae itself, which we refined Zhang et al.'s (2001) definition of to include type species. Therizinosaurids first split into a clade of <i>Erliansaurus</i>, <i>Neimongosaurus</i>, <i>Suzhousaurus</i> and <i>Therizinosaurus</i>. Forcing the former two to be outside a clade of <i>Suzhousaurus</i>, <i>Therizinosaurus </i>and the taxa below, as in Zanno's tree, takes 5 more steps. Notably, we did not include the hindlimb IGM 100/45 in the <i>Therizinosaurus </i>OTU since there's no overlap and its not even particularly large. But here's the <i>Therizinosaurus </i>OTU including the hindlimb. Using this version of <i>Therizinosaurus </i>leaves the tree basically the same but destabilizes it somewhat in that <i>Therizinosaurus </i>and <i>Erlikosaurus </i>can now go in multiple positions within Therizinosauridae, and the Nanchao embryos are in a trichotomy with the <i>Suzhousaurus </i>clade and the <i>Nothronychus </i>clade. Is this an indication the hindlimb produces homoplasy and so might not belong to <i>Therizinosaurus</i>?<br />
<br />
Therizinosaurus ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????01 0110021000 01?0001100 00001110?? ?????????? ?????????? ?????????? ???0???010 1?0000000? 0?1??1???? ?????????? ?????????? 000??0???? ??????3??1 ?????????0 ???00????? 0?1?0?1??? ?????0???? ?????????1 ?????????? ?????0???? ?????????? ?????????? 1011010101 100100??1? ????{12}?00?? 00?12110?? 101??????? 0????0?111 00?010???? ????????1? ?????????? ????100111 1101110??? ?????????? ????????1? ?1???????? ?????????1 ???0?????? ?000?????? ?????0???1 ?????????? ?????????? ?????????? ?????????? ?????0???? ??????-00? ?00-000001 0?0100??0? 10000???0{01} ?{01}?0000??0 {01}0???????? ????????00 ?00?????0- -??11-???? ?????????? ???1?0???0<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVZYtThGu1eES9Z2JKUp-k9nJ3uej0dSjKg0v7ReXsCj-VK_N2IDu0Tu4JG8DAy2qr_jbqKcQo0UMFM40RkLFVkAQEtvKmGjdffdGnrPKtqi_0psP8ygMudQkdaf4gMf7isMvHQiPMK4w/s1600/Segnosaurus+100+83+cerv+neural+arch.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVZYtThGu1eES9Z2JKUp-k9nJ3uej0dSjKg0v7ReXsCj-VK_N2IDu0Tu4JG8DAy2qr_jbqKcQo0UMFM40RkLFVkAQEtvKmGjdffdGnrPKtqi_0psP8ygMudQkdaf4gMf7isMvHQiPMK4w/s320/Segnosaurus+100+83+cerv+neural+arch.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Segnosaurus galbinensis</i> paratype (IGM 100/83) cervical neural arch in right lateral view (courtesy of Zanno).</td></tr>
</tbody></table>
Now comes the Nanchao therizinosaur embryos, those described by Kundrat et al. inside dendroolithid eggs. While including such young specimens might be seen as risky, my ontogenetically conservative scoring method with state N seems to have worked fine here. They fall out where you'd expect a Santonian-Campanian therizinosaur to do so. Following that is <i>Nanshiungosaurus brevispinus</i>, which Senter et al. (2012) recovered as the next most derived therizinosaur after <i>Alxasaurus</i>. Forcing it into this basal position takes 4 steps. <i>Nanshiungosaurus? bohlini </i>was included but pruned a posteriori since it can go anywhere in the <i>Segnosaurus </i>plus <i>Nothronychus </i>clade. Forcing <i>Nanshiungosaurus </i>monophyly is just a single step longer though, while forcing <i>bohlini </i>to be sister to the contemporaneous <i>Suzhousaurus </i>takes 2 steps.<br />
<br />
<i>Segnosaurus </i>itself (which Zanno also provided photos of) pairs with ex-<i>Alectrosaurus </i>forelimb AMNH 6368, which has only previously been analyzed by Zanno (2006) where it pairs with <i>Erliansaurus</i>. Forcing that here compared to other taxa she included results in trees 3 steps longer. <i>Erlikosaurus </i>groups with the <i>Nothronychus </i>species in a trichotomy where it can be sister to either species. Forcing <i>Nothronychus </i>monophyly takes only a single step, but note that no proposed <i>Nothronychus </i>characters involve elements that can be compared to <i>Erlikosaurus </i>(humerus and pes). Forcing <i>Erlikosaurus </i>to group with <i>Therizinosaurus </i>as in Senter et al. requires only a single step, with <i>Erlikosaurus </i>moving to the <i>Therizinosaurus </i>clade.<br />
<br />
Next time, oviraptorosaurs...<br />
<br />
<b>References</b>-
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions
of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.<br />
<br />
Zhang, Xu, Sereno, Kwang and Tan, 2001. A long-necked therizinosauroid
dinosaur from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol, People’s
Republic of China. Vertebrata PalAsiatica. 39(4), 282-290.<br />
<br />
Zanno, 2006. The pectoral girle and forelimb of the primitive therizinosauroid
<i>Falcarius utahensis</i> (Theropoda, Maniraptora): Analyzing evolutionary
trends within Therizinosauroidea. Journal of Vertebrate Paleontology. 26(3),
636-650. <br />
<br />
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of
Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic
Palaeontology. 8(4), 503-543.<br />
<br />
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24(10), 2197-2216.<br />
<br />
<br />
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790. <br />
<br />
Sues and Averianov, 2015. Therizinosauroidea (Dinosauria: Theropoda) from the
Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.<br />
<br />
<span style="color: black;"><span style="font-family: inherit , "times new roman" , serif;">Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Società Paleontologica Italiana. 57(1),
1-25.</span></span><br />
<br />
<span style="color: black;"><span style="font-family: inherit , "times new roman" , serif;">Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. <a href="https://peerj.com/articles/7247" target="_blank">DOI: 10.7717/peerj.7247</a> </span></span> <br />
<br />Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com4tag:blogger.com,1999:blog-3248412803814730250.post-43277181085719446102019-08-19T17:11:00.000-07:002019-08-22T15:50:38.909-07:00Alvarezsaurs in the Lori matrixThis time our topology is-<br />
<div class="separator" style="clear: both; text-align: center;">
<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgAn6MEJmeYTgiqknZl_sPhDdzaMKOBBEBZyF-uYEc5i3Zs7fo8vgELCn2Z9YkPdwLy_vu2khtDiQStxFy5gjU6TlQaCKAUm8Oq6pZPYyFIAyqxTN-d3dTrcQ8VI3iovgtkOpovuU-3V-4/s1600/Alvarez+phylo.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="331" data-original-width="625" height="169" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgAn6MEJmeYTgiqknZl_sPhDdzaMKOBBEBZyF-uYEc5i3Zs7fo8vgELCn2Z9YkPdwLy_vu2khtDiQStxFy5gjU6TlQaCKAUm8Oq6pZPYyFIAyqxTN-d3dTrcQ8VI3iovgtkOpovuU-3V-4/s320/Alvarez+phylo.jpg" width="320" /></a></div>
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I provide a new definition for Alvarezsauroidea that adds <i>Therizinosaurus </i>as an external specifier since I find it most parsimonious for Therizinosauria to be its sister group, and uses <i>Alvarezsaurus </i>as the internal specifier unlike Sereno's that uses <i>Shuvuuia</i>- (<span style="font-style: italic;">Alvarezsaurus calvoi</span> < - <span style="font-style: italic;">Ornithomimus velox, Therizinosaurus cheloniformis, Passer domesticus</span>) . Alvarezsauroids have had a controversial phylogenetic placement, with the Lori matrix recovering them as basal maniraptorans sister to therizinosaurs. Yet they can be outside therizinosaurs plus pennaraptorans in 3 steps, become arctometatarsalians in 4 steps (can bring therizinosaurs or not), non-maniraptoriforms in 6 steps (they bring therizinosaurs), closer to pennaraptorans than therizinosaurs in 6 steps, paravians in 11 steps (therizinosaurs move with), closer to <i>Compsognathus </i>than to birds in 15 steps, closer to birds than deinonychosaurs in 27 steps, and closer to <i>Archaeopteryx </i>and other birds than to dromaeosaurids and troodontids in 30 steps.<br />
<br />
<i>Fukuivenator </i>is an odd taxon, recovered here as the basalmost alvarezsauroid. But it can be a therizinosaurian in only two steps, and outside Maniraptoriformes in 4 steps (it emerges in Coeluridae). One thing I don't think it is is a dromaeosaurid, as that takes 27 more steps, and getting it into Paraves or Pennaraptora requires 11 and 7 steps respectively. Still, I wouldn't be surprised to see this taxon work its way around the base of Maniraptoriformes once an osteology comes out.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjFGe9an-8Tw4Xlrl6gV99Zl6TzBW4UCvJ3hIkFc0MJy5kPTRORUHwNAxSIgysX6OfM7MgVuN1SPI9yoq5geFkeE80thxwCGyI-V0oGxM0lgLmEgS5qO8lWUcW7PuF0vnSVz7NogPfCkcA/s1600/Shuvuuia+975+axial+vent.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjFGe9an-8Tw4Xlrl6gV99Zl6TzBW4UCvJ3hIkFc0MJy5kPTRORUHwNAxSIgysX6OfM7MgVuN1SPI9yoq5geFkeE80thxwCGyI-V0oGxM0lgLmEgS5qO8lWUcW7PuF0vnSVz7NogPfCkcA/s320/Shuvuuia+975+axial+vent.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Shuvuuia deserti</i> IGM 100/975 axial elements in ventral view and pelvis in dorsal view (courtesy AMNH).</td></tr>
</tbody></table>
<i>Nqwebasaurus </i>was recently redescribed by Sereno (2017), which I incorporated into its scorings. Choiniere et al. (2012) recovered it in Ornithomimosauria, but note most of the characters they list to support that are also said to be present in alvarezsauroids. Even they could place it in Alvarezsauroidea with only 4 steps. The Lori matrix needs 6 steps to place it in Ornithomimosauria, which I think is higher partially due to it finding <i>Pelecanimimus </i>to be an alvarezsauroid too. So similarities between the two like their teeth being in a common groove and maxillary teeth being confined to the anterior third of the bone are no longer ornithomimosaur-like. As recently noted by Cerroni et al. (2019), this makes more sense biogeographically as well. Oh, and note that the Lori matrix found <i>Afromimus </i>to be a ceratosaur as in that paper. In any case, <i>Nqwebasaurus </i>takes 10 steps to move to Compsognathidae, and 7 steps to move sister to Pennaraptora.<br />
<br />
As for <i>Pelecanimimus </i>itself, it seems plausibly alvarezsauroid if you think about it. The skull is famously similar to <i>Shuvuuia</i>, the posterior tympanic recess is in the otic recess, ossified sterna are otherwise unknown for ornithomimosaurs, the long manual digit I was always out of place compared to <i>Harpymimus</i>, and Europe makes more sense for otherwise Gondwanan clades in the Cretaceous. Now if only someone would release Perez-Moreno's thesis describing it in detail...<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg0Fh3vp2vKRhFspOnUjH4Plb2vW-tBhKKOQe8jUFVnie9vIYJ1vlARI-an8tTc-6F7iNFTdIt-6-yft3oWlTzBLhhd9kEYSco_0zI080gQF_xF8VmP7w7mV37rztZ-tUjVS_C9LTOmSGk/s1600/Shuvuuia+975+forelimb%257E1.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg0Fh3vp2vKRhFspOnUjH4Plb2vW-tBhKKOQe8jUFVnie9vIYJ1vlARI-an8tTc-6F7iNFTdIt-6-yft3oWlTzBLhhd9kEYSco_0zI080gQF_xF8VmP7w7mV37rztZ-tUjVS_C9LTOmSGk/s320/Shuvuuia+975+forelimb%257E1.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Shuvuuia deserti</i> IGM 100/975 pectoral and forelimb elements. Note the tiny phalanx from digit II or III at the bottom (courtesy AMNH).</td></tr>
</tbody></table>
<i>Patagonykus </i>and <i>Bonapartenykus </i>are usually closer to parvicursorines than <i>Alvarezsaurus </i>and <i>Achillesaurus</i>, but the Lori matrix found them just outside Alvarezsauridae instead. Interestingly, Xu et al. (2018) recovered the same results. It takes 3 steps to move <i>Patagonykus </i>closer to parvicursorines, and 4 steps to join <i>Alvarezsaurus </i>and <i>Patagonykus </i>to the exclusion of parvicursorines as in Alifanov and Barsbold (2009). Xu et al. recover these in 5 and 7 steps respectively, and the
most recent version of Longrich and Currie's alvarezsaurid matrix (Lu et al., 2018) recovers
a basal <span style="font-style: italic;">Patagonykus</span> and a basal Parvicursorinae in 3 steps each. <br />
<br />
One odd result is that the newly described <i>Xiyunykus </i>and <i>Bannykus </i>fall in Patagonykinae too. Yet only 2 steps move them outside the <i>Patagonykus </i>plus Parvicursorinae clade, where they form a clade. Another step breaks that up to place <i>Xiyunykus </i>more basal as in Xu et al.. Them being basal certainly fits better stratigraphically, and Xu et al. use several characters designed for alvarezsauroids that the Lori matrix didn't include yet. Hopefully full osteologies will be published as well.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg81ni3j8QG6FONqWJRgUZBXqYyJRCOVOMxYWLC4bpXDashTloOaZCe6vu3n7WB6MJPj2z-YqCCckIKN4rfWJ9Q_wMVwNkbfFtZRSnsxo_H6Q8L-GoC9s6vQnUvQ_Wapaw-iJgYGm_unVY/s1600/pesplantar.bmp" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="593" data-original-width="600" height="316" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg81ni3j8QG6FONqWJRgUZBXqYyJRCOVOMxYWLC4bpXDashTloOaZCe6vu3n7WB6MJPj2z-YqCCckIKN4rfWJ9Q_wMVwNkbfFtZRSnsxo_H6Q8L-GoC9s6vQnUvQ_Wapaw-iJgYGm_unVY/s320/pesplantar.bmp" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Mononykus olecranus</i> cast YPM 56693 (of holotype) pes in plantar view (courtesy of Senter).</td></tr>
</tbody></table>
A patagonykine <i>Achillesaurus </i>as suggested by Agnolin et al. (2012) takes 7 additional steps in the Lori matrix
where it instead emerges just closer to parvicursorines than <span style="font-style: italic;">Alvarezsaurus</span>. On the other hand, only a single step joins it with <span style="font-style: italic;">Alvarezsaurus</span> as in Longrich and Currie (2009) and only 2 steps makes it just further from parvicursorines than <span style="font-style: italic;">Alvarezsaurus</span> as in Xu et al. (2018).<br />
<br />
<i>Alnashetri </i>is known from type hindlimb material, but now also from MPCA 377, a nearly complete specimen with interesting characters like flat and unfused sternal plates. Makovicky et al. (2016) used this data to recover it as the sister group to Alvarezsauridae, and while the few published details left it more derived in the Lori tree, it can go to a more basal position with only two steps. It should be interesting to compare to e.g. <i>Bannykus </i>once it is published.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhT752fJFzuURtivtQqe89a-T56NTl1tpnwN0gEGv9ORVHjPn_FIiD50CWVrABG26H_SUpQnr3gnZlaVoQrgAqrurKu_m17UAQm0khQ5Lqzgt3prWexq0eDmUV3vWOy5cQ6Zn9uAsVA8Zs/s1600/vertleft.bmp" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1399" data-original-width="1191" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhT752fJFzuURtivtQqe89a-T56NTl1tpnwN0gEGv9ORVHjPn_FIiD50CWVrABG26H_SUpQnr3gnZlaVoQrgAqrurKu_m17UAQm0khQ5Lqzgt3prWexq0eDmUV3vWOy5cQ6Zn9uAsVA8Zs/s320/vertleft.bmp" width="272" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Mononykus olecranus</i> cast YPM 56693 (of holotype) (courtesy of Senter).</td></tr>
</tbody></table>
The arctometatarsal clade has a unique topology, but no other analysis has included nearly as many characters or all of these taxa, with Lu et al. omitting <i>Albinykus </i>and <i>Ceratonykus </i>among non-fragmentary specimens, and Xu et al. omitting the more recently described <i>Qiupanykus</i>. Enforcing the Lori topology in Lu et al.'s matrix is only 5 steps longer, and doing so in Xu et al.'s matrix is only 6 steps longer. On the other hand, Xu et al.'s topology is so unresolved at this level, the only difference in mine is placing the <i>Albinykus </i>plus <i>Xixianykus </i>clade basally near <i>Albertonykus</i>, which takes 5 steps to do in the Lori matrix.<br />
<br />
It should be noted that Lu et al.'s illustrated topology (their Figure 3) is not their matrix's real result, as they did not fully analyze tree space. Instead of 20 trees, there are 214 trees. These differ in that <i>Albertonykus</i>, YPM 1049 and undescribed 41HIII-0104 can fall out anywhere more derived than <i>Patagonykus</i>, and that <i>Parvicursor</i>, the Tugriken Shireh taxon, <i>Shuvuuia </i>and <i>Mononykus </i>form an unresolved polytomy. This leaves <i>Linhenykus</i>, <i>Qiupanykus </i>and <i>Xixianykus </i>unresolved between that polytomy and <i>Patagonykus</i>, which is perfectly compatible with the Lori topology. This may also show that the small alvarezsauroid-specific matrix of Longrich and Currie is insufficient given all the new taxa described since 2009. YPM 1049 was far too fragmentary to include (distal metatarsal III) but I tried testing undescribed Quipa specimen 41HIII-0104. Didn't make it into the publication, but here's its scorings-<br />
<br />
'41HIII0104' ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???????-?? ?????????1 ?10??????? ?????????? ?????????? ???0?????? ????1????? ???1?????{01} ?????????? ?????????? ?????????? ????????0? ????????3? ?????????? ?????????? ?????????? ?????????? ?????????1 ?????????1 1????????? 1????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????0? ?????????? ?????????? ?????????? ?????????? ?????????? ???1?????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????{123}???? ?????????? ?????????? ?????????? ?????????? ?????????? ??1???0???<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEglDikHqzk4qMFebPOS7XWx9Ua7ww91DYHvAnUZ5xzL2h0SekK2PJvQ0JnC_k0EcSzZKE3hKQ2yRhpNcntzCIcNcSVIfWj49136-qQu9d6GQ0MGyffKH0W3KvzpjhpDBagJQeCEhoYEDrI/s1600/Tugrik+99+vent+other.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEglDikHqzk4qMFebPOS7XWx9Ua7ww91DYHvAnUZ5xzL2h0SekK2PJvQ0JnC_k0EcSzZKE3hKQ2yRhpNcntzCIcNcSVIfWj49136-qQu9d6GQ0MGyffKH0W3KvzpjhpDBagJQeCEhoYEDrI/s320/Tugrik+99+vent+other.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in ventral view, forelimb and fibula in lower right (courtesy AMNH).</td></tr>
</tbody></table>
Interestingly, Agnolin et al., Xu et al. and the Lori analysis all recovered <i>Albinykus</i> sister to <span style="font-style: italic;">Xixianykus</span> outside Parvicursorinae. Wonder if that's a real signal? Unfortunately, the only attempt to name this clade was Agnolin et al. who also recovered <i>Ceratonykus </i>in there and called it Ceratonykini. Xu et al. place <i>Ceratonykus </i>closer to parvicursorines, while I found it more basal than either, sister to <i>Qiupanykus </i>which neither of the other studies used. Forcing <i>Ceratonykus </i>sister to <i>Albonykus </i>plus <i>Xixianykus </i>takes 3 more steps in the Lori matrix. Forcing <i>Ceratonykus </i>sister to <i>Mononykus </i>as in its original description (with or without <span style="font-style: italic;">Qiupanykus</span>) takes 5 more steps. As stated in the paper, we were the first analysis to include Hateg tibiotarsi <i>Bradycneme </i>and <i>Heptasteornis</i>. While the former can fall into many positions in Maniraptora, the latter was resolved as an alvarezsaurid as proposed by Naish and Dyke (2004). Note this used only the tibiotarsus and not alvarezsaurid-like distal femur FGGUB R.1957. A single step moves <i>Heptasteornis </i>to Troodontidae.<br />
<br />
We also provide an updated definition for Parvicursorinae <i>(Mononykus olecranus</i> + <i>Parvicursor remotus)</i>, like Choiniere et al.'s (2010) but using species. One accident of our definitional and discovery history is that all these newer arctometatarsal alvarezsaurids (<i>Xixianykus</i>, <i>Albertonykus</i>, <i>Albinykus</i>, <i>Linhenykus</i>, <i>Qiupanykus</i>, <i>Ceratonykus</i>, etc.) emerge outside the originally discovered and defined Parvicursorinae. We could really use some clade defining taxa closer to <i>Mononykus </i>than <i>Patagonykus</i>, <i>Alvarezsaurus </i>or <i>Achillesaurus</i>. In any case, I got a lot of experience with parvicursorine specimens, examining <i>Shuvuuia </i>and the Tugriken Shireh specimen IGM 100/99 in person, and having photos of high quality casts of <i>Mononykus </i>thanks to Senter. I found the Tugriken Shireh taxon closer to <i>Shuvuuia</i>, but moving it closer to <i>Parvicursor </i>as in Longrich and Currie is just 1 step longer.<br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh7CcjmMy_hjhTJooJhCDOx_ShYNHokB94OZxi7vHUqhkXrIh7NSsRG3Ga4wtYjmXv_yDJ-tqf-b0kwYqSbKzDMmXOPmUAG69OArdAh72jG_XjmcYm1rRkvV2U3rTccZO9TQNJynqcAygg/s1600/Tugrik+99+dors+other.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh7CcjmMy_hjhTJooJhCDOx_ShYNHokB94OZxi7vHUqhkXrIh7NSsRG3Ga4wtYjmXv_yDJ-tqf-b0kwYqSbKzDMmXOPmUAG69OArdAh72jG_XjmcYm1rRkvV2U3rTccZO9TQNJynqcAygg/s320/Tugrik+99+dors+other.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Tugriken Shireh parvicursorine (IGM 100/99) vertebrae and ilia in dorsal view, forelimb and fibula in lower right (courtesy AMNH).</td></tr>
</tbody></table>
Next time, therizinosaurs...<br />
<br />
<b>References</b>-
Naish and Dyke, 2004. <i>Heptasteornis</i> was no ornithomimid, troodontid,
dromaeosaurid or owl: The first alvarezsaurid (Dinosauria: Theropoda) from Europe.
Neus Jahrbuch für Geologie und Paläontologie. 7, 385-401. <br />
<br />
Alifanov and Barsbold, 2009. <i>Ceratonykus oculatus</i> gen. et sp. nov., a
new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia.
Paleontological Journal. 43(1), 94-106.<br />
<br />
Longrich and Currie, 2009. <i>Albertonykus borealis</i>, a new alvarezsaur (Dinosauria:
Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for
the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1),
239-252.<br />
<br />
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod
from the early Late Jurassic of Xinjiang, China. Science. 327, 571-574. <br />
<br />
Agnolin, Powell, Novas and Kundrat, 2012. New alvarezsaurid
(Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia
with associated eggs. Cretaceous Research. 35, 33-56.<br />
<br />
Makovicky, Apesteguia and Gianechini, 2016. A new, almost complete specimen of <span style="font-style: italic;">Alnashetri cerropoliciensis</span>
(Dinosauria: Theropoda) impacts our understanding of alvarezsauroid
evolution. XXX Jornadas Argentinas de Paleontologia de Vertebrados.
Libro de resumenes, 74. <br />
<br />
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.<br />
<br />
Lu, Xu, Chang, Jia, Zhang, Gao, Zhang, Zhang and Ding, 2018. A new
alvarezsaurid dinosaur from the Late Cretaceous Qiupa Formation of
Luanchuan, Henan Province, central China. China Geology. 1, 28-35. <br />
<br />
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang,
Xing and Tan, 2018. Two Early Cretaceous fossils document transitional
stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8.
DOI: 10.1016/j.cub.2018.07.057 <br />
<br />
Cerroni, Agnolin, Egli and Novas, 2019. The phylogenetic position of <i>Afromimus tenerensis</i> Sereno, 2017 and its paleobiogeographical implications. Journal of African Earth Sciences. DOI: 10.1016/j.jafrearsci.2019.103572<br />
<br />
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. <a href="https://peerj.com/articles/7247" target="_blank">DOI: 10.7717/peerj.7247</a>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com18tag:blogger.com,1999:blog-3248412803814730250.post-65696267524215823052019-08-10T04:45:00.000-07:002019-08-10T04:45:40.556-07:00Ornithomimosaurs in the Lori matrixNext up is Ornithomimosauria. The <a href="http://theropoddatabase.com/Ornithomimosauria.htm#Ornithomimosauria" target="_blank">Theropod Database has been updated with new ornithomimosaur info</a> too. The published cladogram here is-<br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiBRCWgtzwwzSIwvnZmIOcR4GkmHvW4ZEqbU3S6ER_ORP6Wk2UP2Ewf0HpgGvQtkXKdfoVBVI1rrtR6VJCdgaI7X1ucjBeZoMXmikQs8mLPBH2jAqHiycZpeMSguT8pN_bkivO5Ft1k_7E/s1600/Ornithom0.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="378" data-original-width="556" height="217" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiBRCWgtzwwzSIwvnZmIOcR4GkmHvW4ZEqbU3S6ER_ORP6Wk2UP2Ewf0HpgGvQtkXKdfoVBVI1rrtR6VJCdgaI7X1ucjBeZoMXmikQs8mLPBH2jAqHiycZpeMSguT8pN_bkivO5Ft1k_7E/s320/Ornithom0.jpg" width="320" /></a></div>
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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 <i>Limusaurus </i>or <i>Elaphrosaurus</i>. 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 <i>Deltadromeus</i>. Here's its new scorings<br />
<br />
????????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?<br />
<br />
When Angeac moves in the new matrix with added taxa, we get a little change where <i>Harpymimus </i>is more basal and the polytomy is resolved-<br />
<br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0CuB4dY_st_DtI5L04bhJGokiElF7YH5M4PbkucInHOhKlvazf3gBB7k650C_zt2mKhFNPgzAz1hDRkyy1jK0HyYq8d2FWSlKuO0wVjSlWJZvemJoIjWJBFp3VkKRahp-zAHgltxOMgQ/s1600/Ornithom+taxa+added.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="376" data-original-width="609" height="197" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0CuB4dY_st_DtI5L04bhJGokiElF7YH5M4PbkucInHOhKlvazf3gBB7k650C_zt2mKhFNPgzAz1hDRkyy1jK0HyYq8d2FWSlKuO0wVjSlWJZvemJoIjWJBFp3VkKRahp-zAHgltxOMgQ/s320/Ornithom+taxa+added.jpg" width="320" /></a></div>
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Btw, I published a list of phylogenetic definitions in the supplementary material, including the first to define Ornithomimosauria on the type species- (<i>Ornithomimus velox</i> < - <i>Tyrannosaurus rex, Shuvuuia
deserti, Therizinosaurus cheloniformis, Oviraptor philoceratops, Troodon formosus,
Passer domesticus</i>).<br />
<br />
Ours is one of the few published studies using the new <i>Deinocheirus </i>skeletons, and unlike Lee et al. (2014) I recovered it basal to garudimimids and any other ornithomimosaurs except for <i>Hexing</i>. 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 <i>Sciurumimus </i>section the scoring in Choiniere's analysis is pretty bad.<br />
<br />
This is the first time <i>Hexing </i>has been in an analysis besides the ornithomimosaur-only one in its original description, which did not include <i>Deinocheirus</i>. It groups with <i>Deinocheirus</i>, which is funny because of the size difference, but there would have to be something to fill up that ghost lineage if <i>Deinocheirus </i>is so basal.<br />
<br />
<i>Shenzhousaurus </i>and <i>Harpymimus </i>are next. <i>Kinnareemimus </i>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 <i>Shenzhousaurus </i>and <i>Harpymimus</i>. An extra step moves it to Alvarezsauroidea, so that's still pretty uncertain.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiENp4JMQGfdb9O3ZXbcUhg17E80REr-6ZOirdF9-GDat66QJIYF34XvcveU0ndQUookokHWkMJU5NEQYrEBztcO_yGwkVMEXvk7XeGaAs_Gzad6wVZr6TmxnhifkdobeeiL_wSMDr1gvk/s1600/Archaeornithomimus+AMNH+21800%257E1.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiENp4JMQGfdb9O3ZXbcUhg17E80REr-6ZOirdF9-GDat66QJIYF34XvcveU0ndQUookokHWkMJU5NEQYrEBztcO_yGwkVMEXvk7XeGaAs_Gzad6wVZr6TmxnhifkdobeeiL_wSMDr1gvk/s320/Archaeornithomimus+AMNH+21800%257E1.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Archaeornithomimus asiaticus</i> paratype left femur in posterior view (AMNH 21800) (Courtesy AMNH).</td></tr>
</tbody></table>
<br />
While I didn't recover <i>Deinocheirus </i>by <i>Garudimimus</i>, I did get <i>Beishanlong </i>there as in Lee et al.. Also interesting is that <i>Archaeornithomimus </i>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 <i>Archaeornithomimus bissektensis</i> 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-<br />
<br />
???????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<br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh4oDno7xEMX4WTb1J73ox4gGoQmIOE0EM7PkXZBrdXwls8MAj9SWyWXUwzjBgCUolJSnmKhe7T_sWZUf5QVRyC8iji3Vyv4J1-nijcqZfVCjr5pRY1PT-shZstO9gIHqRdpGO3AOfxxMs/s1600/Archaeornithomimus+bissektensis.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="918" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh4oDno7xEMX4WTb1J73ox4gGoQmIOE0EM7PkXZBrdXwls8MAj9SWyWXUwzjBgCUolJSnmKhe7T_sWZUf5QVRyC8iji3Vyv4J1-nijcqZfVCjr5pRY1PT-shZstO9gIHqRdpGO3AOfxxMs/s320/Archaeornithomimus+bissektensis.jpg" width="183" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Archaeornithomimus? bissektensis</i> holotype right femur (CCMGE 726/12457) (Courtesy of Averianov).</td></tr>
</tbody></table>
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The "Grusimimus" specimen GIN 960910KD ends up in Garudimimidae too, despite generally being seen as a potential juvenile <i>Harpymimus</i>. Joining the two takes 4 steps, so is still quite possible. I wonder if this really is an endemic Asian clade. Interestingly, <i>Arkansaurus </i>(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 <i>Suskityrannus </i>was placed by its authors) with one step. The Lori matrix isn't that great with pedal characters yet.<br />
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We also provide a definition for Ornithomimidae using <i>O. velox</i> and not dependent on the controversial <i>Deinocheirus</i>- (<i>Ornithomimus velox </i> < - <i>Garudimimus brevipes</i>). At the base of Ornithomimidae, we have <i>Nedcolbertia</i>. While an ornithomimosaurian <i>Nedcolbertia</i>'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 <i>Zuolong </i>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 <i>Nedcolbertia</i>'s anatomy, as should the description of BYU 19114 from the Cedar Mountain Formation, said to be similar by Hunt and Quinn. <br />
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Interestingly, <i>Ornithomimus velox</i> does not group with <i>Dromiceiomimus </i>(which includes <i>O. edmontonicus</i> here), but is instead down by <i>Sinornithomimus</i>. The only character really grouping traditional <i>Ornithomimus </i>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 <i>Ornithomimus </i>sensu lato is monophyletic based on the metacarpal length, and I don't think anyone's actually used <i>O. velox</i> as its own OTU before this. It would be funny if <i>Sinornithomimus </i>ended up actually being the Chinese sister taxon to <i>Ornithomimus</i>. <i>Aepyornithomimus </i>is sister to this pair in the published matrix, but in a trichotomy with <i>Ornithomimus</i>, <i>Sinornithomimus </i>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 <i>Archaeornithomimus</i>.<br />
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Next is a novel clade of <i>Tototlmimus </i>and "<i>Gallimimus</i>" "mongoliensis", the first time the latter has been included in an analysis. Only 3 steps are needed to move it sister to <i>Gallimimus bullatus</i>, and <i>Tototlmimus </i>follows. <i>Tototlmimus </i>was poorly resolved in its initial analysis based on Kobayashi characters. Another new clade follows- <i>Rativates </i>plus Kaiparowitz supposed <i>O. velox</i> specimen MNA Pl.1762A plus <i>"Ornithomimus" sedens</i>, the latter based only on the holotype. Again, <i>Rativates </i>has only been analyzed once before in its initial description, using Choiniere's heavily misscored matrix, while <i>sedens</i> and the Kaiparowitz specimen have never been analyzed before. <br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhldsq3z0-TI9cFeEZyEbZDRVKHtXZPT31ZFMeLV7J3HE0VhwAzaB6O3WUqe66SCi_aEscSwDeNc_301clghfF20D3-QWDK732QMP55fq0_j4Y8JfIvwgyMNC5FHdZv7JZrBrDGxgdBilM/s1600/ornithomimosaur+coracoid+comp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="773" data-original-width="1080" height="229" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhldsq3z0-TI9cFeEZyEbZDRVKHtXZPT31ZFMeLV7J3HE0VhwAzaB6O3WUqe66SCi_aEscSwDeNc_301clghfF20D3-QWDK732QMP55fq0_j4Y8JfIvwgyMNC5FHdZv7JZrBrDGxgdBilM/s320/ornithomimosaur+coracoid+comp.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Ornithomimosaur coracoids in right lateral and proximal views. Top left- <i>Beishanlong </i>(after Makovicky et al., 2010). Bottom left- <i>Anserimimus </i>(after Barsbold, 1988). Top right- <i>Sinornithomimus </i>(after Kobayashi, 2004). Bottom right- <i>Gallimimus </i>(after Kobayashi, 2004). Peach dot indicates coracoid tubercle, green line indicates lateral edge of infraglenoid buttress.</td></tr>
</tbody></table>
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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 <i>Anserimimus </i>plus <i>Gallimimus </i>clade, then an American clade of <i>Struthiomimus </i>and '<i>Ornithomimus</i>'. 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 <i>Sinornithomimus</i> and bottom right is <i>Gallimimus</i>. 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 <i>Gallimimus</i>, but compare to the then unknown <i>Beishanlong </i>coracoid in the upper left. It has a hugely protruding process but isn't a part of the <i>Anserimimus </i>plus <i>Gallimimus </i>clade. It's not a commonly shown perspective, but is also found in <i>Nqwebasaurus</i>, <i>Allosaurus</i>, 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 <i>Beishanlong </i>also has this anteriorly positioned, but more problematically <i>Anserimimus </i>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. <i>Adasaurus</i>' pelvis). Note that the more recent Xu et al. (2011) ornithomimosaur analysis that finds the Kobayashi arrangment misscores <i>Beishanlong </i>for both characters. Instead, the Lori analysis (and my previous unpublished TWiG analysis incorporating Kobayashi's and other ornithimomid-relevent characters) recovers a <i>Struthiomimus </i>plus <i>Gallimimus </i>clade and an <i>Anserimimus </i>plus <i>Dromiceiomimus </i>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.<br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj9zjP4jxT1LseNnD4wyiC3n_AhwPV32hgHP3QOTv5zW1654JrMZw_fMpx2l0fwgmD4S5qLgI2rxfIib_Jmg5q37Vsi5dJlZAWXLrFkoUtBdaQiF-i9SJ_bW3RVB8T8VOhHIcHeCvru8Wg/s1600/Timimus+comp.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="870" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj9zjP4jxT1LseNnD4wyiC3n_AhwPV32hgHP3QOTv5zW1654JrMZw_fMpx2l0fwgmD4S5qLgI2rxfIib_Jmg5q37Vsi5dJlZAWXLrFkoUtBdaQiF-i9SJ_bW3RVB8T8VOhHIcHeCvru8Wg/s320/Timimus+comp.jpg" width="173" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Tyrannoraptoran femora in lateral and/or anterior views. Top left- <i>Xiongguanlong </i>(after Li et al., 2010). Top right- <i>Alioramus </i>(after Brusatte et al., 2012). Center- <i>Timimus </i>(after Benson et al., 2012). Bottom left- <i>Archaeornithomimus</i> (courtesy AMNH). Bottom right- <i>Garudimimus </i>in lateral view (after Kobayashi and Barsbold, 2005) and <i>Gallimimus </i>in anterior view (after Osmolska et al., 1972). Peach indicates accessory trochanter in anterior view, green outline indicates accessory trochanter in lateral view.</td></tr>
</tbody></table>
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Also in the <i>Struthiomimus </i>plus <i>Gallimimus </i>group, we get <i>Timimus </i>as the sister taxon to the latter genus. But of course <i>Timimus </i>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 <i>Xiongguanlong </i>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 <i>T. hermani</i> (e.g., <i>Tyrannosaurus</i>)." "<i>T. hermani</i> also possesses a proximomedially inclined (‘elevated’) femoral head, a synapomorphy of derived tyrannosauroids (e.g., <i>Tyrannosaurus</i>; <i>Xiongguanlong</i>), that is absent in ornithomimosaurs." <br />
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First of all, <i>Gallimimus </i>has an elevated femoral head too (Osmolska et al., 1972: Plate XLVII). Not as much as most tyrannosauroids', but neither does <i>Timimus</i>. 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 <i>Xiongguanlong</i>, ~52% in <i>Alioramus</i>), unlike <i>Archaeornithomimus </i>(~69%). But <i>Garudimimus</i>' 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 <i>Garudimimus</i>, 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 <i>Timimus</i>' is more tyrannosauroid in side view, but ironically because they're larger than at least <i>Archaeornithomimus </i>and <i>Gallimimus </i>(green highlight), contra Benson et al.'s statement. And again some ornithomimosaurs like <i>Garudimimus </i>have large accoessory trochanters too. Regarding transverse width, I can't see a difference between e.g. <i>Alioramus </i>and <i>Gallimimus </i>above (peach highlight). I certainly wouldn't say <i>Alioramus</i>' is thicker. So is <i>Timimus </i>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 <i>Aniksosaurus</i>, also Gondwanan Early Cretaceous with a tall and narrow anterior trochanter.<br />
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Finally, <i>Qiupalong </i>joins the <i>Anserimimus </i>plus <i>Dromiceiomimus </i>clade. In its description, it grouped in the American clade, but that's the same Xu et al. (2011) analysis noted above that misscores <i>Beishanlong </i>as lacking the supposed <i>Anserimimus </i>plus <i>Gallimimus </i>characters.<br />
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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 <i>Harpymimus</i>, <i>Beishanlong</i>, <i>Garudimimus</i>, <i>Archaeornithomimus</i>, <i>Sinornithomimus </i>and the derived clade. Overall I'd say this is the best ornithomimosaur analysis published, in taxon number, character number and robusticity of results.<br />
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Next, alvarezsauroids...<br />
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<b>References</b>- Osmólska, Roniewicz and Barsbold, 1972. A new dinosaur, <i>Gallimimus
bullatus</i> n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia.
Palaeontologica Polonica. 27, 103-143.<br />
<br />
Barsbold, 1988. A new Late Cretaceous ornithomimid from the
Mongolia People’s Republic. Journal of Paleontology. 1988(1), 122-125.<br />
<br />
Smith and Galton, 1990. Osteology of <i>Archaeornithomimus asiaticus</i> (Upper
Cretaceous, Iren Dabasu Formation, People's Republic of China). Journal of Vertebrate
Paleontology. 10(2), 255-265.<br />
<br />
Kobayashi, 2004. Asian ornithomimosaurs. PhD thesis. Southern Methodist University.
340 pp.<br />
<br />
Kobayashi and Barsbold, 2005. Reexamination of a primitive ornithomimosaur,
<i>Garudimimus brevipes</i> Barsbold, 1981 (Dinosauria: Theropoda), from the
Late Cretaceous of Mongolia. Canadian Journal of Earth Sciences. 42(9), 1501-1521. <br />
<br />
Li, Norell, Gao, Smith and Makovicky, 2010. A longirostrine tyrannosauroid from
the Early Cretaceous of China. Proceedings of the Royal Society B. 277(1679),
183-190. <br />
<br />
Makovicky, Li, Gao, Lewin, Erickson and Norell, 2010. A giant ornithomimosaur
from the Early Cretaceous of China. Proceedings of the Royal Society B. 277,
191-198. <br />
<br />
Dal Sasso and Maganuco, 2011. <i>Scipionyx samniticus</i> (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Società
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.<br />
<br />
Xu, Kobayashi, Lu, Lee, Liu, Tanaka, Zhang, Jia and Zhang, 2011. A new ornithomimid
dinosaur with North American affinities from the Late Cretaceous Qiupa Formation
in Henan Province of China. Cretaceous Research. 32(2), 213-222.<br />
<br />
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.<br />
<br />
Brusatte, Carr and Norell, 2012. The osteology of <i>Alioramus</i>, 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. <br />
<br />
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body
plan culminated in rapid rates of evolution across the dinosaur-bird
transition. Current Biology. 24(20), 2386-2392. DOI:
10.1016/j.cub.2014.08.034 <br />
<br />
Lee, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and Tsogtbaatar,
2014. Resolving the long-standing enigmas of a giant ornithomimosaur <i>Deinocheirus
mirificus</i>. Nature. 515, 257-260.<br />
<br />
Claessens and Loewen, 2015. A redescription of <i>Ornithomimus velox</i> Marsh,
1890 (Dinosauria, Theropoda). Journal of Vertebrate Paleontology. e1034593 DOI:
10.1080/02724634.2015.1034593 <br />
<br />
Sues and Averianov, 2015. Ornithomimidae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan. Cretaceous Research. 57,
90-110.<br />
<br />
Brownstein, 2017. Redescription of Arundel Clay ornithomimosaur material and a reinterpretation of <span style="font-style: italic;">Nedcolbertia justinhofmanni</span> as an "ostrich dinosaur": Biogeographic implications. PeerJ 5:e3110. DOI: 10.7717/peerj.3110 <br />
<br />
Hunt and Quinn, 2018. A new ornithomimosaur from the Lower Cretaceous
Trinity Group of Arkansas, Journal of Vertebrate Paleontology.
e1421209. DOI: 10.1080/02724634.2017.1421209 <br />
<br />
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. <a href="https://peerj.com/articles/7247/?td=bl" target="_blank">DOI: 10.7717/peerj.7247</a>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com14tag:blogger.com,1999:blog-3248412803814730250.post-55038477281277150662019-08-08T05:54:00.002-07:002019-08-08T05:54:49.593-07:00Basal coelurosaurs in the Lori matrixOkay, I'm on vacation and one of those goals is to get these blog posts out about the results of the <i>Hesperornithoides </i>analysis of Hartman et al. (2019), and the Theropod Database updated again. With that in mind, <a href="http://theropoddatabase.com/Updates.htm" target="_blank">here's our first section</a>.<br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj938VlQY1CFig2gADVcgCG2gCe84XzP9boVz6JJNsb0p9zOKDRaFShhvkNqtXg2_MCeyV7RjIr05Poo1yLe5jd2R4UO44Sqk7WnNAglo2xnmAhSaZsQTR8lbOF6ucT2l7PNjrwDuZEkEc/s1600/Compsog+phylo.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="232" data-original-width="463" height="160" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj938VlQY1CFig2gADVcgCG2gCe84XzP9boVz6JJNsb0p9zOKDRaFShhvkNqtXg2_MCeyV7RjIr05Poo1yLe5jd2R4UO44Sqk7WnNAglo2xnmAhSaZsQTR8lbOF6ucT2l7PNjrwDuZEkEc/s320/Compsog+phylo.jpg" width="320" /></a></div>
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<i>Aristosuchus </i>was analyzed, but is not in the cladogram. We found it to be a coelurosaur closer to Aves than Megaraptora, excluded from Maniraptoriformes. So it could be a basal tyrannosauroid or a compsognathid, or something in between. <br />
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I was able to examine the <i>Ornitholestes </i>holotype at the AMNH twice, and get some nice photos I've been posting here over the years. Besides taking 10 steps to move to Tyrannosauroidea, it takes 7 steps to move outside Tyrannoraptora, 13 steps to move to Maniraptora, and a whopping 30 steps to move to Dromaeosauridae. Should be the nail in the coffin for that idea.<br />
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<i>Sinocalliopteryx </i>emerged basal to compsognathids, but moves to Compsognathidae in 4 steps and joins basal tyrannosauroids in 1 step. Might be worth reexamining its relationship with <i>Dilong</i>.<br />
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<i>Huaxiagnathus </i>and <i>Juravenator </i>join to form a clade basal to compsognathids, but can join that family in a single step. They take 7 steps to move closer to <i>Dilong</i>, so are probably not tyrannosauroids.<br />
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<i>Archaeornithoides </i>is a highly controversial taxon, which I was the first to include in a published analysis. While it most parsimoniously is down here at the compsognathid-grade, only 1 step moves it to Paraves. There it groups with <i>Xiaotingia </i>and <i>Mei </i>as a sinovenatorine instead of by any known Djadochta troodontids. <br />
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<i>Aniksosaurus </i>emerged down here too, more crownward than its original description. There, Martinez and Novas (2006) proposed <span style="font-style: italic;">Aniksosaurus</span>
was a coelurosaur outside Maniraptoriformes, the latter containing
tyrannosaurids in their phylogeny. While not included in a
phylogenetic analysis, they did list three characters supporting this
placement. "Distal tibia with astragalar surface proportionally low"
is also true in basal tyrannosauroids (<span style="font-style: italic;">Coelurus</span>, <span style="font-style: italic;">Guanlong</span>) and maniraptoromorphs (<span style="font-style: italic;">Aorun</span>, <span style="font-style: italic;">Compsognathus</span>, alvarezsauroids including <span style="font-style: italic;">Nqwebasaurus</span>). "Insertion of the M. caudifemoralis longus extensive and deep" is plesiomorphic for maniraptoromorphs, also being found in <span style="font-style: italic;">Ornitholestes</span>, <span style="font-style: italic;">Juravenator</span>
and all ornithomimosaurs. Robust limb elements were stated to resemble
carnosaurs and megalosauroids more than most coelurosaurs, specifically
the humerus, ulna, femur, tibia and pes. The humerus is also robust in
compsognathids and alvarezsaurids, while the metatarsus is less robust
than <span style="font-style: italic;">Ornitholestes</span>, therizinosaurs or <span style="font-style: italic;">Deinocheirus</span>.
However, any measure of basic robusticity such as femoral circumference
should also account for size correlation as larger taxa are typically
less gracile than smaller taxa. Brusatte (2013:395) stated "it is unclear if it possesses any clear
coelurosaurian characters" so did not include it in his TWiG analysis.
However, every analysis including <span style="font-style: italic;">Aniksosaurus</span>
has recovered it within Coelurosauria, and only unquantified
appendicular robusticity has even been suggested to validly support a
more rootward placement than Tyrannoraptora. While I've previously suggested some alvarezsauroid-like characters (e.g. large
presacral neural canals*, ventrally keeled proximal caudal centra,
transversely broad manual ungual I*, laterally expanded brevis shelf*,
distally projecting lateral femoral condyle), enforcing this result in
the Hartman et al. matrix adds 9 steps. However, the asterisked
characters were not included, so it could potentially only be 6 steps
longer. <br />
<br />
<i>Scipionyx </i>gets a pretty traditional position with not much to say.<br />
<br />
<i>Tugulusaurus </i>is a compsognathid here but is historically a very basal coelurosaur, and more recently an alvarezsauroid by Xu et al. (2018). Check <a href="http://theropoddatabase.com/Coelurosauria.htm#Tugulusaurusfaciles" target="_blank">the Database for details</a>, but basically I found basal placements were due to the same three characters, which probably converged in derived tyrannoraptorans and derived maniraptoromorphs. The position in Alvarezsauroidea was based on four characters I didn't examine ("strong medial tab on metacarpal [I]";
"dorsolaterally and dorsomedially facing lateral and medial surfaces of
phalanx [I]-1 that are shallowly concave"; "axial furrow along the
flexor surface of phalanx [I]-1"; "partially enclosed ‘flexor notches’
on the medial and lateral surfaces of the proximal end of the ventral
surface of manual ungual [I]-2"), while it takes 5 more steps to force
as an alvarezsauroid in the Lori matrix. Thus the true difference between a
compsognathid and alvarezsauroid placement could be as low as a single
step.<br />
<br />
<i>Sinosauropteryx </i>is also fairly boring, but photos of NIGP 127587 sent by Senter show that some aspects of its figures in Currie and Chen (2001) aren't preserved in the actual material.<br />
<br />
<i>Santanaraptor </i>gets <a href="http://theropoddatabase.com/Coelurosauria.htm#Santanaraptorplacidus" target="_blank">another long entry</a>, regarding the evidence for tyrannosauroid affinities. Basically almost all of the claimed characters don't exist in <i>Santanaraptor </i>or are impossible to check based on published material and that sent by Bruno Campos. <br />
<br />
<i>Haplocheirus </i>was a surprise as a compsognathid. I know I've seen it outside Alvarezsauroidea in more analyses than Lee and Worthy (2011). Any of you readers remember which? One interesting thing I noticed when moving this entry is that the supposedly autapomorphic dorsally expanded anterior dentary with enlarged teeth is present in all other compsognathids, but the character wasn't included in the Lori analysis. That's always a good sign when it comes to a position's plausibility.<br />
<br />
<i>Compsognathus </i>takes 8 steps to be maniraptoran, 13 to be tyrannosauroid and 15 to be outside Tyrannoraptora. While I did not split the specimens as separate OTUs, they do differ in 16 characters, so I plan to in the future. <br />
<br />
<i>Sciurumimus </i>gets a HUGE section in the supplementary info of the Lori paper. Rauhut asked us to justify its inclusion in the analysis when it's supposed to be a megalosauroid. So I fully reexamined Rauhut et al.''s (2012) analyses of <i>Sciurumimus </i>in the various analyses and found misscorings in each (Smith et al., 2008; Benson et al., 2010; Choiniere et al., 2010) were responsible for keeping it away from <i>Compsognathus</i>. So there's no mystery as to why <i>Juravenator </i>moved to a basal position when <i>Sciurumimus </i>was included but was a coelurosaur when it wasn't, it's just misscorings.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEisznFfNVWf25CqY8GD9F202AbeV0X0L5jsq2fKisJ3Zod9Q1sZU8Eu9aA1teTsrc9R8xZMoX8km6aWXNIY2JFULThKvzwXENk6emGH7LRRoHiuxAntLboUqC8ekMYRMW_aBkhDmSJiC4I/s1600/Sinosauropteryx+skull.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="791" data-original-width="1170" height="216" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEisznFfNVWf25CqY8GD9F202AbeV0X0L5jsq2fKisJ3Zod9Q1sZU8Eu9aA1teTsrc9R8xZMoX8km6aWXNIY2JFULThKvzwXENk6emGH7LRRoHiuxAntLboUqC8ekMYRMW_aBkhDmSJiC4I/s320/Sinosauropteryx+skull.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Skull of NGMC 2124.</td></tr>
</tbody></table>
NGMC 2124 is the <i>Sinosauropteryx </i>sp. of Ji and Ji (1997) and has never been included in an analysis before this. It ends up closer to other compsognathids than <i>Sinosauropteryx</i>, but only takes 3 steps to group with the latter genus. I examined this one in person when it was on display at the RTMP.<br />
<br />
Finally, we have <i>Aorun</i>. Xu et al. (2018) recently proposed this <span style="font-style: italic;"></span>is the basalmost alvarezsauroid based on four unambiguous
synapomorphies- dorsoventrally flattened internarial bar; collateral
ligament fossae on metacarpal I absent; manual digit I bearing large
ungual and all other unguals distinctly smaller; proximodistally
oriented step-like ridge on anterior surface of tibia that braces astragalar ascending process. We recovered <span style="font-style: italic;">Aorun</span>
as a compsognathid, and found it takes 17 more steps to move to
Alvarezsauroidea. This is similar to Cau's (2018) results, where <span style="font-style: italic;">Aorun</span> and <span style="font-style: italic;">Compsognathus</span>
are both non-tyrannoraptoran coelurosaurs and can form a
Compsognathidae with no extra steps, but it takes 9 extra steps to
force an alvarezsauroid <span style="font-style: italic;">Aorun</span>. Notably, even Xu et al.'s matrix only needs 4 steps to move <span style="font-style: italic;">Aorun</span>
to Compsognathidae. Of Xu et al.'s proposed alvarezsauroid
characters, the flattened internarial bar was used by us
and is shared with the controversially compsognathid <span style="font-style: italic;">Haplocheirus</span>. <span style="font-style: italic;">Aorun</span>
actually seems to have a weak medial ligament pit on metacarpal I
(Choiniere et al., 2013: Fig. 15A). The ratio between manual
ungual I and II lengths (1.28) in <span style="font-style: italic;">Aorun</span> is not more than <span style="font-style: italic;">Sciurumimus</span> (~1.29) or <span style="font-style: italic;">Sinosauropteryx</span>
(~1.79). Finally, the ridge bracing the astragalar ascending
process is primitive for maniraptoriforms and also present in <span style="font-style: italic;">Compsognathus</span> and <span style="font-style: italic;">Tugulusaurus</span>. Thus a compsognathid identity is the best supported given proposed characters.<br />
<br />
I was curious with <i>Tugulusaurus</i>, <i>Haplocheirus </i>and <i>Aorun </i>each recovered as alvarezsauroid by Xu et al. and compsognathid by me, is there some unseen connection between the clades? Enforcing them to be more closely related to each other than to other groups takes 15 more steps, so apparently it's not a great hypothesis. <br />
<br />
Next, ornithomimosaurs...<br />
<br />
<b>References</b>- Ji and Ji, 1997. Advances in the study of the avian <i>Sinosauropteryx
prima</i>. Chinese Geology. 242, 30-32.<br />
<br />
Currie and Chen, 2001. Anatomy of <i>Sinosauropteryx prima</i> from Liaoning,
northeastern China. Canadian Journal of Earth Science. 38(12), 1705-1727.<br />
<br />
Martínez and Novas, 2006. <i>Aniksosaurus darwini</i> gen. et sp. nov.,
a new coelurosaurian theropod from the early Late Cretaceous of central Patagonia,
Argentina. Revista del Museo Argentino de Ciencias Naturales. 8(2), 243-259.<br />
<br />
Smith, Makovicky, Agnolin, Ezcurra, Pais and Salisbury, 2008. A <span style="font-style: italic;">Megaraptor</span>-like
theropod (Dinosauria: Tetanurae) in Australia: Support for faunal
exchange across eastern and western Gondwana in the Mid-Cretaceous.
Proceedings of the Royal Society B. 275(1647), 2085-2093. DOI:
10.1098/rspb.2008.0504<br /><br />
Benson, Carrano and Brusatte, 2010. A new clade of archaic large-bodied
predatory dinosaurs (Theropoda: Allosauroidea) that survived to the
latest Mesozoic. Naturwissenschaften. 97(1), 71-78. DOI:
10.1007/s00114-009-0614-x<br /><br />
Choiniere, Clark, Forster and Xu, 2010. A basal coelurosaur
(Dinosauria: Theropoda) from the Late Jurassic (Oxfordian) of the
Shishugou Formation in Wucaiwan, People's Republic of China. Journal of
Vertebrate Paleontology. 30(6), 1773-1796. DOI:
10.1080/02724634.2010.520779 <br />
<br />
Lee and Worthy, 2011. Likelihood reinstates <span style="font-style: italic;">Archaeopteryx</span> as a primitive bird. Biology Letters. 8(2), 299-303. DOI: 10.1098/rsbl.2011.0884<br />
<br />
Rauhut, Foth, Tischlinger and Norell, 2012. Exceptionally preserved juvenile
megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic
of Germany. Proceedings of the National Academy of Sciences. 109(29), 11746-11751.<br />
<br />
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods
(Archosauria: Dinosauria) and patterns of morphological evolution
during the dinosaur-bird transition. PhD thesis, Columbia University.
944 pp.<br />
<br />
Choiniere, Clark, Forster, Norell, Eberth, Erickson, Chu and Xu, 2013 [physically
published 2014]. A juvenile specimen of a new coelurosaur (Dinosauria: Theropoda)
from the Middle-Late Jurassic Shishugou Formation of Xinjiang, People's Republic
of China. Journal of Systematic Palaeontology. 12(2), 177-215.<br />
<br />
<span style="color: black;"><span style="font-family: inherit, Times New Roman, serif;">Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Società Paleontologica Italiana. 57(1),
1-25. DOI: 10.4435/BSPI.2018.01</span></span><br />
<br />
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang,
Xing and Tan, 2018. Two Early Cretaceous fossils document transitional
stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8.
DOI: 10.1016/j.cub.2018.07.057<br />
<br />
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. <a href="https://peerj.com/articles/7247/?td=bl" target="_blank">DOI: 10.7717/peerj.7247</a>Mickey Mortimerhttp://www.blogger.com/profile/08831823442911513851noreply@blogger.com5