Monday, December 26, 2022

Thoughts on Peterson et al. (2022) paper regarding diplodocid feeding mechanisms

Many scientific papers have published regarding the degree of tooth replacement and wear in diplodocoid and macronarian sauropods, namely diplodocids, Camarasaurus, brachiosaurids, and select titanosaurs for which skull material is known. However, very little attention until recently was paid the patterns of tooth replacement and wear in apatosaurine diplodocids, in no small part because the Apatosaurus louisae skull CM 11162, the undescribed Apatosaurus ajax specimen CMC VP-7180, and the apatosaurine specimen TATE 099 found at the Nail Quarry in Como Bluff, Wyoming in 1996, constitute the only apatosaurine skulls that preserve complete teeth (known cranial material for the Apatosaurus ajax holotype includes a braincase and two quadrates, but that's another story). Recently, Peterson et al. (2022) published a paper describing in detail for the first time TATE 099 from Nail Quarry, offering a comprehensive analysis of patterns of tooth wear and replacement for this specimen and further illuminating the nature of feeding mechanisms among flagellicaudatan diplodocoids. 

The history of the systematic placement of TATE 099 within Diplodocidae is rather interesting. It was first referred to as Apatosaurus sp. when first reported in an abstract by Connely and Hawley (1998), who suggested that Apatosaurus used its jaws in front to back sliding motion to aid in cropping and biting when eating plants. For his part, Bakker (1998) referred TATE 099 to Brontosaurus excelsus and used this specimen to claim that B. excelsus differed from Apatosaurus in having the basitubera situated far behind the occipital condyle, but provided no justification as to why TATE 099 was conspecific with B. excelsus, and due to a lack of overlapping material between TATE 099 and B. excelsus holotype, this taxonomic referral was basically ignored by many authors (e.g. Upchurch et al. 2004; Tschopp et al. 2015). In any case, Peterson et al. point out that the basitubera of Apatosaurus louisiae are positioned anterior to the occipital condyle, similar to the condition in TATE 099, and I'm heartened that the authors recognized Bakker's (1998) referral of TATE 099 to Brontosaurus excelsus as lacking basis, and instead assign it to Apatosaurus sp. based on a widely diverging basipterygoid process greater than 60 degrees and the absence of a basisphenoid/basipterygoid recess, both of which are listed as diagnostic characters for Apatosaurus by Tschopp et al. (2015).

CT scans of the maxillae and premaxillae (left) and dentary (right) of Apatosaurus specimen TATE 099 showing unerupted teeth (from Peterson et al. 2022). 3D scans of the unerupted premaxillary/maxillary and dentary teeth shown at the bottom of figures 10 and 11 in Peterson et al. (2022) are included for convenience.

The discussion section of the paper by Peterson et al. (2022) constitutes the real focus on deciphering the rate of tooth wear and replacement among diplodocids. The unerupted tooth counts for TATE 099 reported by the authors in their description of TATE 099 are 5-8 unerupted tooth crowns per alveolus in the premaxilla, 3-5 unerupted tooth crowns per alveolus in the maxilla, and 1-3 unerupted crowns per alveolus in the dentary. Even though Peterson et al. observe differences between diplodocoids and macronarians in the number of replacement teeth as well as tooth volume, shape, and replacement rate, it is quite noteworthy how the authors point out that the rebbachisaurid Nigersaurus lacks alveolar septae and has smaller replacement teeth than those of diplodocid taxa despite the both Nigersaurus and TATE 099 being comparable in the maximum number of replacement teeth. However, the authors don't comment on how the number of replacement teeth of the dentary in TATE 099 compares with that of the holotype of the rebbachisaurid Lavocatisaurus from late Aptian-early Albian deposits in Patagonia, because the latter taxon has several replacement teeth preserved in the dentary. When noting that the numbers of premaxillary/maxillary replacement teeth for TATE 099 as well as Apatosaurus premaxilla MWC 8430 and maxilla MWC 6002 described from Mygatt-Moore Quarry in Colorado by McHugh (2018) are similar, but that dentary of TATE 099 has a number of replacement teeth comparable to those of Dicraeosaurus and the macronarian Brachiosaurus, Peterson et al. (2022) propose that diplodocid taxa retained similar numbers of replacement teeth in the dentary, and interpret the higher numbers of replacement teeth in the upper jaw of Apatosaurus compared to those of diplodocines as supporting the hypothesis by McHugh (2018) that the upper jaw of Apatosaurus was more accustomed to crushing tough plant leaves than that of Diplodocus. In retrospect, the difference between apatosaurines and diplodocines in how they used their upper jaw to chew on plant material may provide another hint at morphological differences between these two diplodocid clades, because some distinguishing features between Apatosaurinae and Diplodocinae can be found in the cervical vertebrae (see Tschopp et al. 2015). Although the erupted tooth row of TATE 099 was unavailable for study by Peterson et al., the authors provide a succinct analysis of the degree of tooth replacement in this specimen based on examination of high-fidelity casts of the erupted tooth row and comparisons with the tooth rows of other sauropods because they note that the row-set tooth replacement patterns in diplodocid taxa like  Apatosaurus, Diplodocus, and Galeamopus contrast with the alternating tooth replacement patterns of the basal macronarian Camarasaurus, since the tooth row of TATE 099 has an uneven occlusal margin that creates an uneven distribution of wear facets on the teeth. 

References:

Bakker, R.T. 1998. Dinosaur mid-life crisis: The Jurassic-Cretaceous transition In Wyoming and Colorado. New Mexico Museum of Natural History and Science Bulletin 14:67-76.

Connely, M.V. and Hawley, R. 1998. A proposed reconstruction of the jaw musculature and other soft cranial tissues of Apatosaurus. Journal of Vertebrate Paleontology 18 (suppl. to volume 3): 35A.

McHugh, J.B. 2018. Evidence for niche partitioning among ground-height browsing sauropods from the Upper Jurassic Morrison Formation of North America. Geology of the Intermountain West 5:95-103.  https://doi.org/10.31711/giw.v5.pp95-103

Peterson, J. E., Lovelace, D., Connely, M., and McHugh, J.B., 2022. A novel feeding mechanism of diplodocid sauropods revealed in an Apatosaurine skull from the Upper Jurassic Nail Quarry (Morrison Formation) at Como Bluff, Wyoming, USA. Palaeontologia Electronica 25(2):a21. https://doi.org/10.26879/1216. palaeo-electronica.org/content/2022/3653-apatosaurine-feeding-mechanism

Tschopp, E., Mateus, O., and Benson, R.B.J. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3:e857.  https://doi.org/10.7717/peerj.857

Upchurch, 
P., 
Tomida
Y., and 
Barrett, 
P.M., 2004. 
A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA
National Science Museum Monographs 
26: 
1
107
.

2 comments:

  1. I began writing this blog in 2013, and I've posted my thoughts on a few papers, including the Mocho et al. (2014) paper about Lourinhasaurus, aspects of the McPhee et al. (2014) paper on Antetonitrus addressing the definition of Sauropodiformes and the affinities of the sauropodiform now known as Kholumolumo (once informally dubbed "Thotobolosaurus"), a rebuttal of the suggestion made by Woodruff and Fowler (2012) that Suuwassea could be a juvenile of a known diplodocid, and my take on the Klamelisaurus paper by Moore et al. (2020).

    Take, for example, my blog post I wrote last year about the Klamelisaurus paper by Moore et al. (2020). In that paper, Euhelopus is recovered in some cladistic analyses outside Macronaria along with Daxiatitan, Dongbeititan, and Xianshanosaurus, but given that Wilson and Upchurch (2009) noted several characters placing Euhelopus in Macronaria, I made the case that Euhelopus could be a basal macronarian because even though it shares some vertebral and dental features with Klamelisaurus, the fact that no caudals are preserved for Euhelopus and the presence of a distolingual boss of the dentition in Yongjinglong led me interpret those characters as having convergently evolved in Euhelopus and Klamelisaurus. The cladistic analyses of Dashanpusaurus and Yuzhoulong with Dai et al. (2022) and Ren et al. (2022) find Euhelopus to be a basal macronarian despite placing this genus outside Somphospondyli, and the recent description of Ruixinia by Mo et al. (2023) shows that derived titanosaurs and select non-titanosaurian macronarians convergently evolved a few vertebral characters with mamenchisaurids, namely the presence of more than 15 cervicals and strongly procoelous anterior and middle caudal vertebrae.

    I decided to write a post for this blog offering a preliminary reappraisal of Bashunosaurus because, when I read the paper by Dai et al. (2022) describing Yuzhoulong, I was caught off guard to see the name Bashunosaurus mentioned in the paper with reference to a hitherto-overlooked paper by Kuang (2004) buried in the references list for the Yuzhoulong paper, given that this name was usually left in the dust as a nomen nudum, and thus I strongly suspected that the paper by Kuang (2004) describes Bashunosaurus as a new taxon, which was confirmed thanks to a PDF of the Kuang (2004) paper sent to me through e-mail by Ren Xinxin (one of the co-authors of the Yuzhoulong paper). In other words, Bashunosaurus was long listed as a nomen nudum in some publications, including Olshevsky (2000), but we now know that it was described as a new taxon in a paper published in 2004.

    Dai, H., Tan, C., Xiong, C., Ma, Q., Li, N., Yu, H., Wei, Z., Wang, P., Yi, J., Wei, G., You, H., and Ren, X., 2022. New macronarian from the Middle Jurassic of Chongqing, China: phylogenetic and biogeographic implications for neosauropod dinosaur evolution. Royal Society Open Science 9 (11). 220794. doi:10.1098/rsos.220794.

    Kuang, X.W., 2004. A new Sauropoda from Kaijiang dinosaur fauna in middle Jurassic beds of North-Eastern Sichuan. pp. 40-46. In: Sun, J.W. (eds), Collection of the 90th anniversary of Tianjin museum of natural history. Tianjin, China: Tianjin Science and Technology Press. (PDF available at http://www.thefossilforum.com/index.php?/topic/127871-bashunosaurus-paper/)

    Mo, J., Ma, F., Yu, Y., and Xu, X., 2023. A new titanosauriform sauropod with an unusual tail from the Lower Cretaceous of northeastern China. Cretaceous Research: in press. doi:10.1016/j.cretres.2022.105449

    Olshevsky, G., 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings 3:1-157.

    Ren, X.X., Jiang, S., Wang, X.R., Peng, G.Z., Ye, Y., King, L., and You, H.L., 2022. Osteology of Dashanpusaurus dongi (Sauropoda: Macronaria) and new evolutionary evidence from Middle Jurassic Chinese sauropods. Journal of Systematic Palaeontology. 20 (1). 2132886. doi:10.1080/14772019.2022.2132886.

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  2. Yes, you've evidently got a lot of solid stuff here. I'm just baffled that I apparently haven't come across it before. Keep up the good work!

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