I am a non-avian dinosaur palaeontologist by trade with a particular interest in their palaeoecology. This can be an endless source of both fascination and frustration. Fascination, because non-avian dinosaurs are quite unlike anything alive today, warranting some use of creative license when imagining them as living animals. Frustration, because the lack of good, extant ecological analogues frequently makes reconstruction of their ancient ecologies an almost insurmountable challenge.

The Canadian Museum of Nature where I work has a good collection of Late Cretaceous turtles. I took an interest in these some years ago because it struck me that, despite the quality of our collection, relatively few people come to study them. I thought, "Someone should work on these. Why not me?" I figured studying a new fossil group would present a fun change of pace and perhaps a more straightforward object of palaeoecological reconstruction. After all, fossil turtles are a lot like living turtles, so how hard can it be? Right?

In 2018, I took a special interest in one recently prepared fossil turtle, which I determined to be a new species of Basilemys (Mallon and Brinkman, 2018). Basilemys held my interest because, although it is a relatively common form, there has been some debate concerning the palaeohabitat of this animal and its closest relatives, the nanhsiungchelyids. Some have argued for an aquatic habitat for these animals; others, for a terrestrial one. It seems that where one comes down on the issue depends on which aspect of ecomorphology is emphasized. If it is on the flat carapace, nanhsiungchelyids must have been aquatic; if it is on the stout feet, terrestrial. This is how I came to appreciate the numerous ecomorphological proxies (e.g., skull shape, shell shape, limb proportions) that are used in turtle palaeoecology and how incongruent they can sometimes be. So much for easy answers!

The present study by Evers et al. is a response to an original piece of research by Lichtig and Lucas (2017), who claimed to be able to use simple shell measurements (carapacial doming and relative plastral width) to accurately deduce/infer the habitats of living turtles and, by extension, fossil ones. In short, they found that terrestrial turtles tend to have more domed carapaces and wider plastra, yielding some unconventional palaeoecological reconstructions of particular stem turtles. Evers et al. take issue with several aspects of this study, including issues of faulty data entry, inappropriate removal of extant taxa from the model, and insufficient accounting for phylogenetic non-independence. By correcting for these overights, they find that the model of Lichtig and Lucas (2017) performs more poorly than advertised and that the palaeoecological classification it produces should be questioned. "The map is not the territory", as Alfred Korzybski put it, and this latest study by Evers et al. serves as an important reminder of that lesson.
Still, even if Lichtig and Lucas's model is overly simplistic, it is true that aquatic turtles, on average, have lower carapaces and narrower plastra, and that they have relatively lower skulls and longer toes. Surely, there is merit in each of these anatomical proxies, even if no single one predicts ecology with total accuracy. I would love to see a model that combines them all. Until then, Evers et al. have inched us closer to knowing what turtle morphology can (and cannot) tell us about habitat.

Thanks to D. Brinkman and H. Smith for their helpful reviews of the manuscript.

References

Evers, S. W., Foth, C., Joyce, W. G., and Hermanson, G. (2024). Simple shell measurements do not consistently predict habitat in turtles: A reply to Lichtig and Lucas (2017). bioRxiv, 586561, ver. 3 peer-reviewed by PCI Paleo. https://doi.org/10.1101/2024.03.25.586561

Lichtig, A. J., and Lucas, S. G. (2017). A simple method for inferring habitats of extinct turtles. Palaeoworld, 26(3), 581–588. https://doi.org/10.1016/j.palwor.2017.02.001

Mallon, J. C., and Brinkman, D. B. (2018). Basilemys morrinensis, a new species of nanhsiungchelyid turtle from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. Journal of Vertebrate Paleontology, 38(2), e1431922. https://doi.org/10.1080/02724634.2018.1431922

Bedane et al. (2024) provide a beautifully illustrated demonstration of the difficulties in using dental adaptation as proxies for the diets of elephants, which are in turn often used to determine the vegetation in an area. This study set out to assess mesowear, which is the relief on the teeth that forms due to abrasion by food, and therefore a good proxy of dietary composition in herbivores. The team was interested in testing whether this mesowear relates to morphological adaptations of hypsodonty (high-crownedness) and enamel thickness over a period of ~3.4–~1.1 million years in an elephant species (Elephas recki) commonly found in the Plio/Pleistocene of the Shungura formation (Omo Valley, Ethiopia). To answer this question, the team scored these metrics in 140 molars between ~3.4 and ~1.1 million years of age, separated into time bins. Their results show surprisingly low levels of variation in mesowear, indicating relatively low variation in diet that was overall mostly composed of graze (as opposed to mixed or browsing diets, which are softer). Hypsodonty and enamel thickness were correlated, but changed erratically rather than suggesting a trend towards a particular dietary adaptation. The exciting conclusion is that dental morphologies that we often consider to be adaptive to certain conditions are very slow to evolve, and that a wide variety of morphologies can support the survival of a species despite little variation in diet. For me as a functional evolutionary morphologist, this clear case of many-to-one-mapping is a timely reminder that evolution does not work either quickly or just on the one character complex I might be considering. And in terms of using elephant teeth as ecological proxies – this job clearly just got a little harder.

References

Bedane, T. G., Mackaye, H. T., and Boisserie, J.-R. (2025). New data on morphological evolution and dietary adaptations of Elephas recki from the Plio-Pleistocene Shungura Formation (Lower Omo Valley, Ethiopia). PaleorXiv, qexuf, ver. 4, peer-reviewed by PCI Paleo. https://doi.org/10.31233/osf.io/qexuf

Fossils get around. Any one fossil locality might be sampled by several collectors from as many institutions around the world. Alternatively, a single collector might heavily sample a site, and sell or trade parts of their collection to other institutions, scattering the fossils far and wide. These practices have the advantage of making fossils from any one locality available to researchers across the globe. However, they also have the disadvantage that, in order to systematically survey any one species, a researcher must follow innumerable trails of breadcrumb to get to where the relevant materials are held.

This is true of many famous fossil localities, such as the Eocene Green River Formation in the USA, the Cretaceous Kem Kem beds of Morocco, or the Devonian Miguasha cliffs of Canada. It is especially true of the Upper Jurassic deposits of the Morrison Formation in the western USA, which have yielded an impressive assemblage of megaherbivorous sauropod dinosaurs over the last 150 years. Today, these bones are to be found in museums not just in the USA, but also in Canada, Argentina, Japan, Australia, Malaysia, South Africa, and throughout Europe. Trawling museum databases in search of sauropod material from the Morrison Formation can therefore be a daunting task, never mind traveling the globe to actually study them.

A new paper by Tschopp et al. (2019) seeks to ease the burden on sauropod researchers by introducing a database of Morrison Formation sauropods, consisting of over 3000 specimens housed in nearly 40 institutions around the world. The authors are themselves sauropod workers and, having suffered first-hand the plight of studying material from the Morrison Formation, came up with a solution to the problem of keeping track of it all. The database is founded largely on material personally seen by the authors, supplemented by information from the literature and museum catalogs. The database further provides information on bone representation, ontogeny, locality details, and fine-scale stratigraphy, among other fields. Like any database, it is a living document that will continue to grow as new finds are made. Tschopp et al. (2019) have wisely chosen to allow others to contribute to the listing, but changes must first be vetted for accuracy. This product represents 10 years of work, and I have little doubt that it will be well-received by those of us who work on dinosaurs. Speaking personally, my PhD research on megaherbivorous dinosaurs from the Dinosaur Park Formation of Canada led me to institutions in Canada, the USA, and the UK, and further stops to Spain and Argentina would have been beneficial, if affordable. Planning for this work would have been greatly assisted by a database like the one provided us by Tschopp et al. (2019). Many a future graduate student will undoubtedly owe them a debt of gratitude.

References

Tschopp, E., Whitlock, J. A., Woodruff, D. C., Foster, J. R., Lei, R., & Giovanardi, S. (2019). The Morrison Formation Sauropod Consensus: A freely accessible online spreadsheet of collected sauropod specimens, their housing institutions, contents, references, localities, and other potentially useful information. PaleorXiv, version 3, peer-reviewed by PCI Paleo. doi: 10.31233/osf.io/ydvra

Mesozoic fishes are extremely diverse. In fact, fishes are the most diverse group of vertebrates during the Mesozoic─just as during any other era. Yet, their study is severely underrepresented in comparison to other fossil groups. There are just too few palaeoichthyologists to deal with such a vast diversity of fishes. Nonetheless, thanks to the huge efforts they have made over the last few decades, we have come a long way in our understanding of Mesozoic ichthyofaunas. One of such devoted palaeoichthyologists is Dr. Adriana López-Arbarello, whose contributions have been crucial in elucidating the phylogenetic interrelationships and taxonomic diversity of Mesozoic actinopterygian fishes (e.g., López-Arbarello, 2012; López-Arbarello & Sferco, 2018; López-Arbarello & Ebert, 2023). In her most recent manuscript, Dr. López-Arbarello has joined forces with Dr. Rainer Brocke to tackle the taxonomy and systematics of the halecomorph fishes from one of the most relevant Triassic sites, the upper Ladinian Perledo locality from Italy (López-Arbarello & Brocke, 2024). 

Fossil fishes were reported for the first time from Perledo in the first half of the 19th century (Balsamo-Crivelli, 1839), and up to 30 different species were described from the locality in the subsequent decades. Unfortunately, this is one of the multiple examples of fossil collections that suffered the effects of World War II, and most of the type material was lost. As a consequence, many of those 30 species that have been described over the years are in need of a revision. Based on the study of additional material that was transferred to Germany and is housed at the Senckenberg Research Institute and Natural History Museum, López-Arbarello & Brocke (2024) confirm the presence of four different species of halecomorph fishes in Perledo, which were previously put under synonymy (Lombardo, 2001). They provide new detailed information on the anatomy of two of those species, together with their respective diagnoses. But more importantly, they carry out a thorough exercise of taxonomy, rigorously applying the International Code of Zoological Nomenclature to disentangle the intricacies in the taxonomic story of the species placed in the genus Allolepidotus. As a result, they propose the presence of the species A. ruppelii, which they propose to be the type species for that genus (instead of A. bellottii, which they transfer to the genus Eoeugnathus). They also propose a new genus for the other species originally included in Allolepidotus, A. nothosomoides. Finally, they provide a set of measurements and ratios for Pholidophorus oblongus and Pholidophorus curionii, the other two species previously put in synonymy with A. bellottii, to demonstrate their validity as different species. However, due to the loss of the type material, the authors propose that these two species remain as nomina dubia. 

In summary, apart from providing new detailed anatomical descriptions of two species and solving some long-standing issues with the taxonomy of the halecomorphs from the relevant Triassic Perledo locality, the paper by López-Arbarello & Rainer (2024) highlights three important topics for the study of the fossil record: 1) we should never forget that world-scale problems, such as World Wars, also affect our capacity to understand the natural world in which we live, and the whole society should be aware if this; 2) the importance of exhaustively following the International Code of Zoological Nomenclature when describing new species; and 3) we are in need of new palaeoichthyologists to, in Dr. López-Arbarello’s own words, “unveil the mysteries of those marvellous Mesozoic ichthyofaunas.”

References

Balsamo-Crivelli, G. (1839). Descrizione di un nuovo rettile fossile, della famiglia dei Paleosauri, e di due pesci fossili, trovati nel calcare nero, sopra Varenna sul lago di Como, dal nobile sig. Ludovico Trotti, con alcune riflessioni geologiche. Il politecnico repertorio mensile di studj applicati alla prosperita e coltura sociale, 1, 421–431.

Lombardo, C. (2001). Actinopterygians from the Middle Triassic of northern Italy and Canton Ticino (Switzerland): Anatomical descriptions and nomenclatural problems. Rivista Italiana di Paleontologia e Stratigrafia, 107, 345–369. https://doi.org/10.13130/2039-4942/5439

López-Arbarello, A. (2012). Phylogenetic interrelationships of ginglymodian fishes (Actinopterygii: Neopterygii). PLOS ONE, 7(7), e39370. https://doi.org/10.1371/journal.pone.0039370

López-Arbarello, A., and Brocke, R. (2024). New generic name for a small Triassic ray-finned fish from Perledo (Italy). PaleorXiv, bxmg5, ver. 4, peer-reviewed by PCI Paleo. https://doi.org/10.31233/osf.io/bxmg5

López-Arbarello, A., and Ebert, M. (2023). Taxonomic status of the caturid genera (Halecomorphi, Caturidae) and their Late Jurassic species. Royal Society Open Science, 10(1), 221318. https://doi.org/10.1098/rsos.221318

López-Arbarello, A., and Sferco, E. (2018). Neopterygian phylogeny: The merger assay. Royal Society Open Science, 5(3), 172337. https://doi.org/10.1098/rsos.172337

There are less and less experts on taxonomy of particular groups particularly among early career paleontologists and (paleo)biologists – this also includes ammonoid cephalopods. Techniques cannot replace this taxonomic expertise (Engel et al. 2021) but machine learning approaches can make taxonomy more efficient, reproducible as well as passing it over more sustainable. Initially ammonoid taxonomy was a black box with small differences sometimes sufficient to erect different species as well as really idiosyncratic groupings of superficially similar specimens (see De Baets et al. 2015 for a review). In the meantime, scientists have embraced more quantitative assessments of conch shape and morphology more generally (see Klug et al. 2015 for a more recent review). The approaches still rely on important but time-intensive collection work and seeing through daisy chains of more or less accessible papers and monographs without really knowing how these approaches perform (other than expert opinion). In addition, younger scientists are usually trained by more experienced scientists, but this practice is becoming more and more difficult which makes it difficult to resolve the taxonomic gap. This relates to the fact that less and less experienced researchers with this kind of expertise get employed as well as graduate students or postdocs choosing different research or job avenues after their initial training effectively leading to a leaky pipeline and taxonomic impediment.

Robust taxonomy and stratigraphy is the basis for all other studies we do as paleontologists/paleobiologists so Foxon (2021) represents the first step to use supervised and unsupervised machine-learning approaches and test their efficiency on ammonoid conch properties. This pilot study demonstrates that machine learning approaches can be reasonably accurate (60-70%) in identifying ammonoid species (Foxon, 2021) – at least similar to that in other mollusk taxa (e.g., Klinkenbuß et al. 2020) - and might also be interesting to assist in cases where more traditional methods are not feasible. Novel approaches might even allow to further approve the accuracy as has been demonstrated for other research objects like pollen (Romero et al. 2020). Further applying of machine learning approaches on larger datasets and additional morphological features (e.g., suture line) are now necessary in order to test and improve the robustness of these approaches for ammonoids as well as test their performance more broadly within paleontology.

References

De Baets K, Bert D, Hoffmann R, Monnet C, Yacobucci M, and Klug C (2015). Ammonoid intraspecific variability. In: Ammonoid Paleobiology: From anatomy to ecology. Ed. by Klug C, Korn D, De Baets K, Kruta I, and Mapes R. Vol. 43. Topics in Geobiology. Dordrecht: Springer, pp. 359–426.

Engel MS, Ceríaco LMP, Daniel GM, Dellapé PM, Löbl I, Marinov M, Reis RE, Young MT, Dubois A, Agarwal I, Lehmann A. P, Alvarado M, Alvarez N, Andreone F, Araujo-Vieira K, Ascher JS, Baêta D, Baldo D, Bandeira SA, Barden P, Barrasso DA, Bendifallah L, Bockmann FA, Böhme W, Borkent A, Brandão CRF, Busack SD, Bybee SM, Channing A, Chatzimanolis S, Christenhusz MJM, Crisci JV, D’elía G, Da Costa LM, Davis SR, De Lucena CAS, Deuve T, Fernandes Elizalde S, Faivovich J, Farooq H, Ferguson AW, Gippoliti S, Gonçalves FMP, Gonzalez VH, Greenbaum E, Hinojosa-Díaz IA, Ineich I, Jiang J, Kahono S, Kury AB, Lucinda PHF, Lynch JD, Malécot V, Marques MP, Marris JWM, Mckellar RC, Mendes LF, Nihei SS, Nishikawa K, Ohler A, Orrico VGD, Ota H, Paiva J, Parrinha D, Pauwels OSG, Pereyra MO, Pestana LB, Pinheiro PDP, Prendini L, Prokop J, Rasmussen C, Rödel MO, Rodrigues MT, Rodríguez SM, Salatnaya H, Sampaio Í, Sánchez-García A, Shebl MA, Santos BS, Solórzano-Kraemer MM, Sousa ACA, Stoev P, Teta P, Trape JF, Dos Santos CVD, Vasudevan K, Vink CJ, Vogel G, Wagner P, Wappler T, Ware JL, Wedmann S, and Zacharie CK (2021). The taxonomic impediment: a shortage of taxonomists, not the lack of technical approaches. Zoological Journal of the Linnean Society 193, 381–387. doi: 10. 1093/zoolinnean/zlab072

Foxon F (2021). Ammonoid taxonomy with supervised and unsupervised machine learning algorithms. PaleorXiv ewkx9, ver. 3, peer-reviewed by PCI Paleo. doi: 10.31233/osf.io/ewkx9

Klinkenbuß D, Metz O, Reichert J, Hauffe T, Neubauer TA, Wesselingh FP, and Wilke T (2020). Performance of 3D morphological methods in the machine learning assisted classification of closely related fossil bivalve species of the genus Dreissena. Malacologia 63, 95. doi: 10.4002/040.063.0109

Klug C, Korn D, Landman NH, Tanabe K, De Baets K, and Naglik C (2015). Ammonoid conchs. In: Ammonoid Paleobiology: From anatomy to ecology. Ed. by Klug C, Korn D, De Baets K, Kruta I, and Mapes RH. Vol. 43. Dordrecht: Springer, pp. 3–24.

Romero IC, Kong S, Fowlkes CC, Jaramillo C, Urban MA, Oboh-Ikuenobe F, D’Apolito C, and Punyasena SW (2020). Improving the taxonomy of fossil pollen using convolutional neural networks and superresolution microscopy. Proceedings of the National Academy of Sciences 117, 28496–28505. doi: 10.1073/pnas.2007324117