A fundamental question in evolutionary biology and paleobiology is how quickly populations and/or species evolve and under what circumstances. Because the fossil record affords us the most direct view of how species lineages have changed in the past, considerable effort has gone into developing methodological approaches for assessing rates of evolution as well as what has been termed “mode of evolution” which generally describes pattern of evolution, for example whether the morphological change captured in fossil time series are best characterized as static, punctuated, or trending (e.g., Sheets and Mitchell, 2001; Hunt, 2006, 2008; Voje et al., 2018). The rock record from which these samples are taken, however, is incomplete, due to spatially and temporally heterogenous sediment deposition and erosion. The resulting structure of the stratigraphic record may confound the direct application of evolutionary models to fossil time series, most of which come from single localities. This pressing issue is tackled in a new study entitled “Identification of the mode of evolution in incomplete carbonate successions” (Hohmann et al., 2024).

The “carbonate successions” part of the title is important. Previous similar work (Hannisdal, 2006) used models for siliciclastic depositional systems to simulate the rock record. Here, the authors simulate sediment deposition across a carbonate platform, a system that has been treated as fundamentally different from siliciclastic settings, from both the point of view of geology (see Wagoner et al., 1990; Schlager, 2005) and ecology (Hopkins et al., 2014). The authors do find that stratigraphic structure impacts the identification of mode of evolution but not necessarily in the way one might expect; specifically, it is less important how much time is represented compared to the size and distribution of gaps, regardless of where you are sampling along the platform. This result provides an important guiding principle for selecting fossil time series for future investigations. 

Another very useful result of this study is the impact of time series length, which in this case should be understood as the density of sampling over a particular time interval. Counterintuitively, the probability of selecting the data-generating model as the best model decreases with increased length. The authors propose several explanations for this, all of which should inspire further work. There are also many other variables that could be explored in the simulations of the carbonate models as well as the fossil time series. For example, the authors chose to minimize within-sample variation in order to avoid conflating variability with evolutionary trends. But greater variance also potentially impacts model selection results and underlies questions about how variation relates to evolvability and the potential for directional change. 

Lastly, readers of Hohmann et al. (2024) are encouraged to also peruse the reviews and author replies associated with the PCI Paleo peer review process. The discussion contained in these documents touch on several important topics, including model performance and model selection, the nature of nested model systems, and the potential of the forwarding modeling approach.  

References

Hannisdal, B. (2006). Phenotypic evolution in the fossil record: Numerical experiments. The Journal of Geology, 114(2), 133–153. https://doi.org/10.1086/499569

Hohmann, N., Koelewijn, J. R., Burgess, P., and Jarochowska, E. (2024). Identification of the mode of evolution in incomplete carbonate successions. bioRxiv, 572098, ver. 4 peer-reviewed by PCI Paleo. https://doi.org/10.1101/2023.12.18.572098

Hopkins, M. J., Simpson, C., and Kiessling, W. (2014). Differential niche dynamics among major marine invertebrate clades. Ecology Letters, 17(3), 314–323. https://doi.org/10.1111/ele.12232

Hunt, G. (2006). Fitting and comparing models of phyletic evolution: Random walks and beyond. Paleobiology, 32(4), 578–601. https://doi.org/10.1666/05070.1

Hunt, G. (2008). Gradual or pulsed evolution: When should punctuational explanations be preferred? Paleobiology, 34(3), 360–377. https://doi.org/10.1666/07073.1

Schlager, W. (Ed.). (2005). Carbonate Sedimentology and Sequence Stratigraphy. SEPM Concepts in Sedimentology and Paleontology No. 8. https://doi.org/10.2110/csp.05.08

Sheets, H. D., and Mitchell, C. E. (2001). Why the null matters: Statistical tests, random walks and evolution. Genetica, 112, 105–125. https://doi.org/10.1023/A:1013308409951

Voje, K. L., Starrfelt, J., and Liow, L. H. (2018). Model adequacy and microevolutionary explanations for stasis in the fossil record. The American Naturalist, 191(4), 509–523. https://doi.org/10.1086/696265

Wagoner, J. C. V., Mitchum, R. M., Campion, K. M., and Rahmanian, V. D. (1990). Siliciclastic Sequence Stratigraphy in Well Logs, Cores, and Outcrops: Concepts for High-Resolution Correlation of Time and Facies. AAPG Methods in Exploration Series No. 7. https://doi.org/10.1306/Mth7510

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

A newly discovered fossil of a tuna fish from the Miocene has been identified in the Duho Formation in Pohang City, South Korea (Suh et al., 2025). The new find, attributed to the genus Auxis, represents only the second valid fossil record of this genus globally, thus contributing to the understanding of evolutionary history within the Scombridae family (Collette and Nauen, 1983; Nam et al., 2021). 

The partially skeleton GNUE322001 consists of a few articulated caudal vertebrae preserving diagnostic features of the genus Auxis (Suh et al., 2025). Although it is not possible to compare the new find with the only fossil species known to date, †A. koreanus Nam et al., 2021, the significant difference in size suggests that it could be a different species. The fossil, preserved in fine-grained mudstone, also offers insights into taphonomic processes, suggesting that the specimen underwent significant decomposition in a low-energy sedimentary environment before burial.

The new record of Auxis supports interpretations of the Duho Formation as a pelagic and subtropical marine habitat, shaped by upwelling activities during the Miocene (Graham and Dickson, 2000; Kim and Paik, 2013; Nam et al., 2021). This discovery emphasizes the significance of upwelling zones in fostering biodiversity and highlights the value of fossil records in reconstructing prehistoric marine ecosystems (Lalli and Parsons, 1997; Wang and Lee, 2019).

References

Collette, B. B., and Nauen, C. E. (1983). Scombrids of the world: An annotated and illustrated catalogue of tunas, mackerels, bonitos, and related species known to date. Food and Agriculture Organization of the United Nations.

Graham, J. B., and Dickson, K. A. (2000). The evolution of thunniform locomotion and heat conservation in scombrid fishes: New insights based on the morphology of Allothunnus fallai. Zoological Journal of the Linnean Society, 129(4), 419–466. https://doi.org/10.1111/j.1096-3642.2000.tb00612.x

Kim, J., and Paik, I. S. (2013). Chondrites from the Duho Formation (Miocene) in the Yeonil Group, Pohang Basin, Korea: Occurrences and paleoenvironmental implications. Journal of the Geological Society of Korea, 49(3), 407–416. https://doi.org/10.14770/jgsk.2013.49.3.407

Lalli, C. M., and Parsons, T. R. (1997). Biological oceanography: An introduction (2nd ed). Butterworth Heinemann.

Nam, G.-S., Nazarkin, M. V., and Bannikov, A. F. (2021). First discovery of the genus Auxis (Actinopterygii: Scombridae) in the Neogene of South Korea. Bollettino Della Società Paleontologica Italiana, 60(1), 61–67. https://doi.org/10.4435/BSPI.2021.05

Suh, D., Kim, S.-H., and Nam, G.-S. (2025). A new tuna specimen (Genus Auxis) from the Duho Formation (Miocene) of South Korea. bioRxiv, 605724, ver. 5 peer-reviewed by PCI Paleo. https://doi.org/10.1101/2024.07.29.605724

Wang, Y.-C., and Lee, M.-A. (2019). Composition and distribution of fish larvae surrounding the upwelling zone in the waters of northeastern Taiwan in summer. Journal of Marine Science and Technology, 27(5), article 8. https://doi.org/10.6119/JMST.201910_27(5).0008