Neimengornis the chimaera, "Yuanchuavis" is another Pengornis and more - October 2021 Database update

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.

The coelophysoid Pendraig was added.  If anyone has Warrener's 1983 thesis originally featuring the material, feel free to send a copy over.  I also researched the nomen nudum Sinosaurus "shawanensis" that was previously represented by one DML post by Olshevsky in 2002, based on my hard copy of The Jurassic System of China-

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 "Sinosaurus shawanensis (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 shawanensis by Young was transferred to Sinosaurus by someone else, but while Young did name Sinosaurus triassicus neither he nor anyone else named a vertebrate species shawanensis (the only animals with that species name before 1985 are brachiopod Cryptospirifer shawanensis Jing et al., 1974 and small shelly fossil Phyllochites shawanensis Duan, 1983, neither from the Lufeng Formation).  Young (1951) did refer one tooth from Shawan to Sinosaurus (IVPP V279), but this was to S. triassicus and was from the Dull Purplish Beds, so doesn't match stratigraphically with Cheng's taxon.  As the holotype and most paratypes of S. triassicus 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 S. triassicus is not listed by Cheng either.  Thus "shawanensis" is near certainly a typo for triassicus, but is still listed here as this can probably not be proven more than thirty years after the fact with the author dead. 
Note Molina-Perez and Larramendi (2019) represent Sinosaurus "shawanensis" with isolated dorsal vertebra IVPP V31, which was referred to Sinosaurus triassicus by Young (1948).  Yet its size and morphology are similar to mid dorsals of sauropodomorph skeleton IVPP V100, also referred to S. triassicus 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 Sinosaurus triassicus", being from the Dark Red Beds as well, and has no connection to the name "shawanensis."

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 Ceratosuchops and Riparovenator.  As a lumper my initial hunch was that they were individual variations of Baryonyx walkeri, but Barker et al. (2021) do an impressive job detailing all of the differences and similarities between these and 'Suchomimus' (Cristatusaurus on my site).  Based on the known anatomical data I don't think we can either synonymize the Wessex taxa or sink them into Baryonyx 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 Cristatusaurus and Suchomimus resemble each other more than Baryonyx - a less rounded anterodorsal margin to their premaxillae (also in Ceratosuchops but not Riparovenator); a comparatively larger second alveolus (also in Riparovenator but not Ceratosuchops).

I also finally moved Camarillasaurus 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, Mahakala 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.

Maniraptoran anterior dorsal (MPG-KPC24) referred to Camarillasaurus cirugedae 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.

Moving to paravians, I added Papiliovenator and also scored it in the Lori matrix where it emerged sister to Zanabazar.  Forcing it to be sister to Linhevenator or Philovenator 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..  

Holotype of Neimengornis rectusmim (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.

Neimengornis 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 Sapeornis, although the skull, tail and furcula are Shenzhouraptor-grade, while the sacrum has an expanded but distally unfused last rib as in Confuciusornis 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 Sapeornis, 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 Dalianraptor where III-1 is), pointing to it being an artificial articulation of elements.  Thus we have a situation much like Dalianraptor.

Snout tip of Yuanchuavis kompsosoura (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).

Wang et al. (2021) described a supposed new taxon of pengornithid- Yuanchuavis kompsosaura 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 Pengornis, IVPP V18632 and Parapengornis (right premaxilla; first tooth displaced and loose in left element).  The supposedly short anterior and posterior lacrimal processes are similar to Parapengornis, IVPP V18632 and Pengornis 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 Pengornis 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, Parapengornis and Chiappeavis.  Finally, the short anterodorsal pygostyle processes are also present in Parapengornis and Chiappeavis (eroded in Pengornis), while the elongate anteroventral process is also present in Parapengornis, partly exposed in Pengornis and probably hidden under caudal vertebrae in Chiappeavis.  Of the variable characters noted above for Parapengornis and Chaippeavis, 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, Yuanchuavis is synonymized with Pengornis here.

Tail comparison of Pengornis 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.

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.

References- Young, 1948. On two new saurischians from Lufeng, Yunnan. Bulletin of the Geological Society of China. 28, 75-90.

Young, 1951. The Lufeng saurischian fauna in China. Palaeontologica Sinica. C(13), 1-96.

Cheng, 1985. The Lufeng-Dafang Subregion. In Wang, Cheng and Wang (eds.). The Jurassic System of China. Stratigraphy of China. 11, 185-189.

Olshevsky, DML 2002. https://web.archive.org/web/20181215172409/http://dml.cmnh.org/2002Apr/msg00630.html

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.

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.

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.

Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.

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.

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 

Samathi, Sander and Chanthasit, 2021 online. A spinosaurid from Thailand (Sao Khua Formation, Early Cretaceous) and a reassessment of Camarillasaurus cirugedae from the Early Cretaceous of Spain. Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.1874372

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. 

Wang, O'Connor, Zhao, Pan, Zheng, Wang and Zhou, 2021. An Early Cretaceous enantiornithine bird with a pintail. Current Biology. 31, 1-8.

Identity of the mysterious "Luckyraptor" plus other new birds

Last year I noticed an entry for "Luckyraptor [Anonymous] 2007 [nomen nudum]" on Olshevsky's Dinosaur Genera List, 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 an old DML post 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.

As you can see, "Luckyraptor eastensis" is based on the Jixiangornis orientalis 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.

 

Harris mentioned three other theropod nomina nuda in Zhang's book, one of which is Sinornithosaurus "zhaoi".  Turns out that's based on QM V1002 that was later briefly described as a Microraptor gui specimen by Xing et al. (2013).  I translate Zhang's Chinese text as-

"Sinornithosaurus zhaoi (New Species)

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 Sinornithosaurus millenii, 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.

Place of Origin: Beipiao, Liaoning

Age: Late Jurassic to Early Cretaceous"

However, its humerofemoral ratio (~86%) is actually shorter than the S. millenii holotype (~91%) and Microraptor teeth are typically less elongate than Sinornithosaurus'.  The slender ischium, short tibia, elongate pedal phalanx IV-4 and long leg remiges all support assigning QM V1002 to Microraptor instead of Sinornithosaurus and it is here considered a junior synonym of M. zhaoianus.  "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).

Next we have "Jinzhouornis" "delicates".  My translation is-

"Jinzhouornis delicates (New Species)

The primitive bird whose individual is smaller than the petite Liaoxiornis, 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 Lycoptera fossils on the slab of Jinzhouornis delicates. 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.

Place of Origin: Yixian County, Liaoning

Age: Early Cretaceous"

The generic assignment is odd because Jinzhouornis is a confuciusornithid (and indeed, a junior synonym of Confuciusornis), 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. Liaoxiornis, Dalingheornis, GMV-2158 and 2159).

Finally, there's "Smallornis liaoningica", whose entry I translate as-

"Smallornis liaoningica (new species)  

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. The forelimbs are covered with very delicate feathers, and the down feathers on the head are more beautifully decorated. Smallornis liaoningica is a very small arboreal bird.  

Place of Origin: Yixian County, Liaoning  

Age: Early Cretaceous"

 

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.

If anyone has further information like specimen numbers, provenance or higher quality scans for "delicates" or "Smallornis", feel free to share.

References- Zhang, 2007. The Fossils of China. China University of Geosciences Press. 502 pp.

Xing, Persons, Bell, Xu, Zhang, Miyashita, Wang and Currie, 2013. Piscivory in the feathered dinosaur Microraptor. Evolution. 67(8), 2441-2445.

"Megalosaurus" cloacinus and more - September 2021 Database Update

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.

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 Megalosaurus species.  silesiacus is a generic carnivorous archosauriform tooth too early to be dinosaurian, while cloacinus 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.

"Zanclodon" silesiacus Jaekel, 1910
= Megalosaurus silesiacus (Jaekel, 1910) Kuhn, 1965
Early Anisian, Middle Triassic
Lower Gogolin Formation, Lower Muschelkalk, Poland
Holotype- (University of Griefswalden/Göttinger coll.; lost?) tooth (24x12x5 mm)
Referred- ?(Geological Museum of the Polish Geological Institute-National Research Institute coll.) tooth (Skawiński, Ziegler, Czepiński, Szermański, Tałanda, Surmik and Niedźwiedzki, 2017)
?(Silesian University of Technology, Faculty of Mining and Geology coll.) tooth (37 mm) (Surmik and Brachaniec, 2013)
Comments- Jaekel (1910) noted (translated) "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, Zanclodon silesiacus. The only difference between [phytosaur Mesorhinosuchus] 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 Megalosaurus.  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."  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 Megalosaurus silesiacus as other material of Zanclodon silesiacus.  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 Teyujagua 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 Megalosaurus or another neotheropod, and the presence of serrations is unlike Zanclodon, 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.
References- Jaekel, 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. 5, 197-229.
Kuhn, 1965. Fossilium Catalogus 1: Animalia. Pars 109: Saurischia. Ysel Press. 94 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Surmik and Brachaniec, 2013. The large superpredators' teeth from Middle Triassic of Poland. Contemporary Trends in Geoscience. 2, 91-94.
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.

 

Holotype tooth of "Zanclodon" silesiacus (University of Griefswalden/Göttinger coll.; lost?) in labial (A), basal section (B) and more apical section (C) (after Jaekel, 1910).

"Megalosaurus" cloacinus Quenstedt, 1858
= Plateosaurus cloacinus (Quenstedt, 1858) Huene, 1905
= Gresslyosaurus cloacinus (Quenstedt, 1858) Huene, 1932
= Pachysaurus cloacinus (Quenstedt, 1858) Huene, 1932
Rhaetian, Late Triassic
Exter Formation, Germany
Syntypes- (lost) two teeth
Referred- ?(GPIT and SMNS coll.) many teeth (Huene, 1905)
?(SMNS 52457) tooth (~25x11x? mm) (Huene, 1905)
?(SMNS coll.) teeth (Roemer, 1870)
? seven teeth (Miller Endlich, 1870)
Norian-Rhaetian?, Late Triassic
'Lisów Breccia', Poland
?(University of Wroclaw coll.; lost) two teeth (Roemer, 1870)
Early Hettangian, Early Jurassic
Calcaire de Valognes, Manche, France
?(University of Caen coll.; destroyed) tooth (Rioult, 1978)
Comments- 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 Lucianosaurus 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) "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 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. Nicrosaurus), and it could even be a fish tooth which often have these types of root expansion.  Huene (1905) listed the species as "Plateosaurus" cloacinus within Theropoda, stating it includes Rhaetian dentary "Zanclodon cambrensis".  In 1908 he places it in Plateosauridae within Theropoda and states (translated) "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."  He illustrated a tooth in figure  274 as "From the Rhaetian Bonebed of Bebenhausen near Tübingen. Tooth in nat. Size. The tip is missing. Original in the natural history cabinet in Stuttgart."  Regarding cambrensis, Huene states "The teeth have the greatest resemblance to Plateosaurus cloacinus 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.  Huene later (1932) assigns cloacinus to Teratosauridae within Carnosauria, listed as both Pachysaurus cloacinus (pg. 6) and Gresslyosaurus cloacinus (pg. 72, 114).  Steel (1970) calls it Gresslyosaurus cloacinus within Plateosauridae.  Buffetaut et al. (1991) mentions "A tooth referred to Megalosaurus cloacinus 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 cloacinus, when Huene stated it was only one of "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, Daemonosaurus), so is here placed in Archosauriformes indet..  Note Huene (1905, 1908) used "Plateosaurus" as a placeholder genus because until 1911 Plateosaurus was thought to have carnivorous teeth, and used "Pachysaurus" and "Gressylosaurus" 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 Megalosaurus teeth are more similar to cloacinus' syntypes and SMNS 52457 than prosauropod teeth.
References- Quenstedt, 1858. Der Jura. H. Laupp'schen. 842 pp.
Miller Endlich, 1870. Das Bonebed Württembergs. Druck Von Ludwig Friedrich Fues. 30 pp.
Roemer, 1870. Geologie von Oberschlesien. Robert Nischkowsky. 587 pp.
Huene, 1905. Uber die Trias-Dinosaurier Europas. Zeitschrift der Deutschen Geologischen Gesellschaft. 57, 345-349.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie. Gustav Fischer Verlag. 1-87.
Rioult, 1978. Inventaire des dinosauriens mésozoïques de Normandie. Ecosystèmes continentaux mésozoiques de Normandie (Livret-guide). Université de Caen. 26-29.
Buffetaut, Cuny and le Loeuff, 1991. French dinosaurs: The best record in Europe? Modern Geology. 16, 17-42.
Dzik and Sulej, 2007. A review of the early Late Triassic Krasiejów biota from Silesia, Poland. Palaeontologia Polonica. 64, 1-27.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
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.

Syntype teeth of "Megalosaurus" cloacinus (lost) in side and sectional views (after Quenstedt, 1858).

Other named specimens from the paper include the archosauromorph femur Zanclodon? "antiquus" based on a museum label, and neotheropod fibula Velocipes.  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 Camposaurus.  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.

The other thing about Skawiński et al.'s Velocipes 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 Coelophysis bauri 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.

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 Spinosuchus (that is seemingly a Trilophosaurus species), and revising the Protoavis 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 Megalancosaurus, 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.

Protoavis texensis holotype proximal femur (TTU-P9200; top) in anterior view (after Chatterjee, 1991) compared to "Megapnosaurus" kayentakatae holotype (MNA.V.2623; bottom) (after Rowe, 1989). Note the muscle scars in the former typical of robust coelophysoids.

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 Saltriovenator paper changing to coelophysoid Dilophosaurus 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 Dilophosaurus in Coelophysoidea, so that would be interesting to compare to Tykoski's (2005) thesis that took twenty additional steps to place Dilophosaurus closer to averostrans.  As part of the coelophysoid revision, Sarcosaurus was updated after Ezcurra et al. (2021).

New abelisaurid Kurupi was added.  An upcoming goal is to change Abelisauridae to the stem (away from Noasaurus and Ceratosaurus), and probably move Noasauridae outside Neoceratosauria.

New tetanurines are Ulughbegsaurus and "Cryptotyrannus", 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.

I also added new unenlagiines Ypupiara and "Lopasaurus".  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 Ypupiara, but unlike the latter is from a different locality than "Lopasaurus".

Finally, we have new birds that were added to the cladograms but not given entries yet.  Youornis is uncontroversial, but Yuanchuavis looks to just be another Pengornis but with typically dimorphic elongate retrices unlike the 'Chiappeavis' specimen.  Wasaibpanchi 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.

See you in a month.

Additional References-  Rowe, 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. Journal of Vertebrate Paleontology. 9(2), 125-136.

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.

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.

Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.

Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.

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. 

Ezcurra and Brusatte, 2011. Taxonomic and phylogenetic reassessment of the early neotheropod dinosaur Camposaurus arizonensis from the Late Triassic of North America. Palaeontology. 54(4), 763-772.

Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2016. Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.

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.