Kjetil Lysne Voje

• Thaureau, Marion & Voje, Kjetil Lysne (2026). Modeling within-lineage evolution as a consequence of changes in the adaptive landscape across short and long timescales. Evolution. ISSN 0014-3820. 80(9), p. 2098–2098. doi: 10.1093/evolut/qpag127. Full text in Research Archive Show summary

  Abstract The adaptive landscape has been suggested as a potential conceptual bridge between phenotypic evolution on generational and macroevolutionary timescales. However, this potential remains largely untapped due to a limited understanding of how it changes through time. Here, we assessed the dynamics of the adaptive landscape across different time intervals by analyzing phenotypic time series using multivariate models of adaptation. First, we examined whether a decrease in river water flow affected the optimal body mass of a salmon population over a few decades. Second, we explored whether temperature changes affected the optimal size of coccoliths in a species of coccolithophore across about a hundred thousand years in the Cretaceous. Finally, we analyzed the extent to which temperature changes affected the optimal shell size in a lineage of planktic foraminifera over a few million years during the Miocene. We find evidence that evolution induced by changes in the adaptive landscape, driven by environmental change, occurred in the salmon and foraminifer datasets, but we do not detect a causal link between the investigated paleo–proxies and size changes in the coccolithophore lineage. We argue that applying the same modeling framework to datasets covering different time spans not only offers a promising way to better connect evolutionary processes across timescales but also enables tests of adaptive hypotheses about within-lineage evolution.

• Pienaar, Jason; Bartoszek, Krzysztof; Brahmantio, Bayu; Fierst, Janna L; Fuentes-González, Jesualdo A & Hansen, Thomas Fredrik [Show all 9 contributors for this article] (2025). Phylogenetic comparative methods for studying adaptation: the adaptation-inertia framework. Journal of Evolutionary Biology. ISSN 1010-061X. 39 (2026)(1), p. 17–17. doi: 10.1093/jeb/voaf113. Full text in Research Archive Show summary

  Abstract Phylogenetic comparative methods are a major tool for evaluating macroevolutionary hypotheses. Methods based on the mean-reverting stochastic Ornstein–Uhlenbeck process allow for modelling adaptation on a phenotypic adaptive landscape that itself evolves and where fitness peaks depend on measured characteristics of the external environment and/or other organismal traits. Here, we give an overview of the conceptual framework for the many implementations of these methods and discuss how we might interpret estimated parameters. We emphasize that the ability to model a changing adaptive landscape sets these methods apart from other approaches and discuss why this aspect captures long-term trait evolution more realistically. Recent multivariate extensions of these methods provide a powerful framework for testing evolutionary hypotheses but are also more complicated to use and interpret. We provide some guidance on their usage and put recent literature on the topic in biological rather than mathematical terms. We further show how these methods provide a starting point for modelling reciprocal selection (i.e., coevolution) between interacting lineages. We then briefly review some critiques of the methodologies. Finally, we provide some ideas for future developments that we think will be useful to evolutionary biologists.

• Pélabon, Christophe; Agudelo-Cantero, Gustavo Adolfo; Ajoy, Yimen Gerardo Araya; Bolstad, Geir Hysing; Cheng, Changde & Galis, Frietson [Show all 26 contributors for this article] (2025). Evolvability: progress and key questions. BioScience. ISSN 0006-3568. 75(12), p. 1057–1057. doi: 10.1093/biosci/biaf111. Full text in Research Archive Show summary

  Since the 1990s, evolutionary biologists have recognized the importance of explaining the ability of biological systems to evolve and how this ability itself evolves. This recognition of the need to explain evolvability emerged from an awareness that the kind and the amount of heritable variation available for natural selection require explanation. The concept of evolvability is now the focus of many research programs in diverse subdisciplines within evolutionary biology. In the present article, we first review and synthesise progress made in evolvability research. We then present key questions to set an agenda for future research on evolvability, identify challenges to answer these questions, and discuss opportunities to apply results from the evolvability research to conservation biology.

• Kinneberg, Vilde Bruhn & Voje, Kjetil L. (2025). Rate–time scaling in phenotypic evolution: Limitations of current models in capturing temporal dynamics. Evolution. ISSN 0014-3820. doi: 10.1093/evolut/qpaf208. Full text in Research Archive Show summary

  Evolutionary rates correlate negatively with time, which makes it complicated to compare rates across lineages that have diversified on different time intervals. The causes of this correlation are debated. Using simulations, we first show that rates of evolution estimated as a parameter in the unbiased random walk model lack a rate–time scaling when data has been generated using this model, even when time series are made incomplete and biased. This indicates that it is theoretically possible to estimate rates that are not time correlated from empirical data. We then analyze 643 empirical time series to assess whether accounting for model misspecification, sampling error, and model identifiability reduces the negative scaling, but none appear to have a significant impact. This suggests that the rate–time correlation requires an explanation grounded in evolutionary biology and that common models used in phylogenetic comparative studies and phenotypic time series analyses often fail to accurately describe trait evolution in empirical data. Making meaningful comparisons of estimated rates between clades and lineages covering different time intervals remains a challenge.

• Schendel, Vanessa; Hamilton, Brett R.; Robinson, Samuel D.; Green, Kathryn; Sayre, Marcel E. & Brown, Darren [Show all 13 contributors for this article] (2025). Exaptation of an evolutionary constraint enables behavioural control over the composition of secreted venom in a giant centipede. Nature Ecology and Evolution. 383. doi: 10.1038/s41559-024-02556-9. Full text in Research Archive Show summary

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• Hunt, Gene; Voje, Kjetil L. & Liow, Lee Hsiang (2025). Punctuated equilibrium: State of the evidence. Paleobiology. ISSN 0094-8373. 51(4), p. 652–661. doi: 10.1017/pab.2024.31. Full text in Research Archive Show summary

  Despite its many extensions and implications, we argue that punctuated equilibrium itself has two core, empirical claims: (1) stasis dominates within fossil species; and (2) morphological change is concentrated in pulses that occur associated with speciation. Here we assess the state of the evidence for these two claims, 50 years after punctuated equilibrium’s foundational paper. Spurred by controversy, paleontologists have amassed a large number of case studies in which morphology in species-level lineages is tracked over time. Modern, likelihood-based methods have been used to fit to these data models of stasis, random walks, and directional trends, as well as more complex dynamics. Compilations reveal that the directional trends predicted by gradualist expectations are infrequent. Although stasis is commonly observed, it is favored in less than half of cases, and meandering random walks or more complex models generally account for the majority of cases. The second claim of punctuated equilibrium has received much less empirical scrutiny than the first. Although speciational pulses are plausible in theory, only a few paleontological studies integrate ancestor–descendant time series into a phylogenetic framework as is needed to estimate cladogenetic change and compare it with anagenesis. These studies, as well as more indirect analyses of extant clades, suggest that speciational change can occur, but we cannot yet assess with confidence its frequency or importance compared with anagenetic changes.

• Holstad, Agnes; Voje, Kjetil L.; Opedal, Øystein Hjorthol; Bolstad, Geir Hysing; Bourg, Salome Cassandra Sarah & Hansen, Thomas Fredrik [Show all 7 contributors for this article] (2024). Evolvability predicts macroevolution under fluctuating selection. Science. ISSN 0036-8075. 384(6696), p. 688–693. doi: 10.1126/science.adi8722. Full text in Research Archive Show summary

  Heritable variation is a prerequisite for evolutionary change, but the relevance of genetic constraints on macroevolutionary timescales is debated. By using two datasets on fossil and contemporary taxa, we show that evolutionary divergence among populations, and to a lesser extent among species, increases with microevolutionary evolvability. We evaluate and reject several hypotheses to explain this relationship and propose that an effect of evolvability on population and species divergence can be explained by the influence of genetic constraints on the ability of populations to track rapid, stationary environmental fluctuations.

• Bartoszek, Krzysztof; Tredgett Clarke, John; Fuentes-González, Jesualdo; Mitov, Venelin; Pienaar, Jason & Piwczyński, Marcin [Show all 9 contributors for this article] (2024). Fast mvSLOUCH: Multivariate Ornstein–Uhlenbeck-based models of trait evolution on large phylogenies. Methods in Ecology and Evolution. 15(9), p. 1507–1515. doi: 10.1111/2041-210X.14376. Full text in Research Archive Show summary

  1. The PCMBase R package is a powerful computational tool that enables efficient calculations of likelihoods for a wide range of phylogenetic Gaussian models. 2. Taking advantage of it, we redesigned the R package mvSLOUCH. 3. Here, we demonstrate how the new version of the package can be used to thoroughly examine the evolution and adaptation of traits in a large dataset of 1252 vascular plants through the use of multivariate Ornstein–Uhlenbeck processes. 4. The results of our analysis demonstrate the ability of the modelling framework to distinguish between various alternative hypotheses regarding the evolution of functional traits in angiosperms.

• Tsuboi, Masahito; Sztepanacz, Jacqueline; Lisle, Stephen De; Voje, Kjetil L.; Grabowski, Mark & Hopkins, Melanie J. [Show all 25 contributors for this article] (2024). The paradox of predictability provides a bridge between micro- and macroevolution. Journal of Evolutionary Biology. ISSN 1010-061X. 37(12), p. 1413–1432. doi: 10.1093/jeb/voae103. Full text in Research Archive Show summary

  The relationship between the evolutionary dynamics observed in contemporary populations (microevolution) and evolution on timescales of millions of years (macroevolution) has been a topic of considerable debate. Historically, this debate centers on inconsistencies between microevolutionary processes and macroevolutionary patterns. Here, we characterize a striking exception: emerging evidence indicates that standing variation in contemporary populations and macroevolutionary rates of phenotypic divergence is often positively correlated. This apparent consistency between micro- and macroevolution is paradoxical because it contradicts our previous understanding of phenotypic evolution and is so far unexplained. Here, we explore the prospects for bridging evolutionary timescales through an examination of this “paradox of predictability.” We begin by explaining why the divergence–variance correlation is a paradox, followed by data analysis to show that the correlation is a general phenomenon across a broad range of temporal scales, from a few generations to tens of millions of years. Then we review complementary approaches from quantitative genetics, comparative morphology, evo-devo, and paleontology to argue that they can help to address the paradox from the shared vantage point of recent work on evolvability. In conclusion, we recommend a methodological orientation that combines different kinds of short-term and long-term data using multiple analytical frameworks in an interdisciplinary research program. Such a program will increase our general understanding of how evolution works within and across timescales.

• Voje, Kjetil L.; Saito-Kato, Megumi & Spanbauer, Trisha L. (2024). Evolution in fossil time series reconciles observations in micro-and macroevolution. Journal of Evolutionary Biology. ISSN 1010-061X. 37(12), p. 1551–1562. doi: 10.1093/jeb/voae087. Full text in Research Archive Show summary

  Extrapolating microevolutionary models does not always provide satisfactory explanations for phenotypic diversification on million-year time scales. For example, short-term evolutionary change is often modelled assuming a fixed adaptive landscape, but macroevolutionary changes are likely to involve changes in the adaptive landscape itself. A better understanding of how the adaptive landscape changes across different time intervals and how these changes cause populations to evolve has the potential to narrow the gap between micro- and macroevolution. Here, we analyze two fossil diatom time series of exceptional quality and resolution covering time intervals of a few hundred thousand years using models that account for different behaviours of the adaptive landscape. We find that one of the lineages evolves on a randomly and continuously changing landscape, whereas the other lineage evolves on a landscape that shows a rapid shift in the position of the adaptive peak of a magnitude that is typically associated with species-level differentiation. This suggests phenotypic evolution beyond generational timescales may be a consequence of both gradual and sudden repositioning of adaptive peaks. Both lineages show rapid and erratic evolutionary change and are constantly readapting towards the optimal trait state, observations that align with evolutionary dynamics commonly observed in contemporary populations. The inferred trait evolution over a span of a few hundred thousand years in these two lineages is, therefore, chimeric in the sense that it combines components of trait evolution typically observed on both short and long timescales.

• Lifjeld, Jan Terje; Cramer, Becky; Leder, Erica Helen & Voje, Kjetil L. (2024). Sperm as a speciation phenotype in promiscuous songbirds. Evolution. ISSN 0014-3820. 79(1), p. 134–143. doi: 10.1093/evolut/qpae154. Full text in Research Archive Show summary

  Sperm morphology varies considerably among species. Sperm traits may contribute to speciation if they diverge fast in allopatry and cause conspecific sperm precedence upon secondary contact. However, their role in driving prezygotic isolation has been poorly investigated. Here we test the hypothesis that, early in the speciation process, female promiscuity promotes a reduction in overlap in sperm length distributions among songbird populations. We assembled a data set of 20 pairs of populations with known sperm length distributions, a published estimate of divergence time, and an index of female promiscuity derived from extrapair paternity rates or relative testis size. We found that sperm length distributions diverged more rapidly in more promiscuous species. Faster divergence between sperm length distributions was caused by the lower variance in the trait in more promiscuous species, and not by faster divergence of the mean sperm lengths. The reduced variance is presumably due to stronger stabilizing selection on sperm length mediated by sperm competition. If divergent sperm length optima in allopatry causes conspecific sperm precedence in sympatry, which remains to be shown empirically, female promiscuity may promote prezygotic isolation and rapid speciation in songbirds.

• Voje, Kjetil L.; Grabowski, Mark; Holstad, Agnes; Porto, Arthur; Tsuboi, Masahito & Bolstad, Geir Hysing (2023). Does lack of evolvability constrain adaptation? If so, on what time scales? In Hansen, Thomas Fredrik; Houle, David; Pavlicev, Mihaela & Pelabon, Christophe (Ed.), Evolvability A Unifying Concept in Evolutionary Biology?. MIT Press. ISSN 9780262545624. p. 289–306. doi: 10.7551/mitpress/14126.001.0001. Full text in Research Archive Show summary

  The relevance of genetic constraints for evolutionary change beyond microevolutionary timescales is debated. The high evolvability of natural populations predicts rapid adaptation, but evolvability is often found to correlate with phenotypic divergence on longer timescales, which makes sense if evolvability constraints divergence. This chapter attempts to reconcile the observation of high evolvability of ppulations with the idea that genetig constraints may still be relevant on long timescales. We first establish that a relationship between evolvability and divergence is a common empirical phenomenon but among populations within species (microevolution) and among species (macroevolution), We then argue that a satisfactory model for the prevalence of this empirical relationship is lacking. Linking microevolutionary theory of phenotypic change on macroevolution timescales - is key to better understanding the relative importance of genetic constraints on phenotypic evolution beyond a handful generations.

• Johannessen, Lars Erik; Johnsen, Arild; Koppetsch, Thore; Lifjeld, Jan Terje; Matschiner, Michael & Søli, Geir Einar Ellefsen [Show all 7 contributors for this article] (2023). Arctic Specimens in the Zoological Collections at the Natural History Museum, University of Oslo, Norway (NHMO). Collections: A Journal for Museum and Archives Professionals. ISSN 1550-1906. doi: 10.1177/15501906231167574. Full text in Research Archive Show summary

  In the NHMO zoological collections, specimens from the Arctic include about 9,000 mammals and 7,100 birds, whereas the Insect Collection holds about 105,000 specimens plus more than hundred jars with unsorted material. The Fish Collection contains approximately 1,400 specimens, while the Herptile Collection (amphibians & reptiles) holds only thirty-one specimens of three taxa. Many of these specimens originate from expeditions to E Greenland, N Canada, Svalbard, Novaya Zemlya, Finnmark, and NE Siberia in the period 1898 to 1966. Furthermore, the DNA Bank has about 5,600 tissue and extracted DNA samples, mostly sampled from wild animals during the last decades but also from specimens in the voucher collections. Most of the Arctic specimens have been digitized and are available in online data portals like GBIF, except for the Insect Collection, where only the type material and about 30 percent of the total specimens are digitized.

• Voje, Kjetil L.; Saulsbury, James Gabriel; Starrfelt, Jostein; Latorre, Daniel Varajão; Rojas, Alexis & Kinneberg, Vilde Bruhn [Show all 10 contributors for this article] (2023). Measurement theory and paleobiology. Trends in Ecology & Evolution. ISSN 0169-5347. 38, p. 1165–1176. doi: 10.1016/j.tree.2023.08.005. Full text in Research Archive Show summary

  Measurement theory, a branch of applied mathematics, offers guiding principles for extracting meaning from empirical observations and is applicable to any science involving measurements. Measurement theory is highly relevant in paleobiology because statistical approaches assuming ratio-scaled variables are commonly used on data belonging to nominal and ordinal scale types. We provide an informal introduction to representational measurement theory and argue for its importance in robust scientific inquiry. Although measurement theory is widely applicable in paleobiology research, we use the study of disparity to illustrate measurement theoretical challenges in the quantitative study of the fossil record. Respecting the inherent properties of different measurements enables meaningful inferences about evolutionary and ecological processes from paleontological data.

• Grabowski, Mark Walter; Pienaar, Jason; Voje, Kjetil L.; Andersson, Staffan; Fuentes-González, Jesualdo & Kopperud, Bjørn Tore [Show all 10 contributors for this article] (2023). A Cautionary Note on "A Cautionary Note on the Use of Ornstein Uhlenbeck Models in Macroevolutionary Studies". Systematic Biology. ISSN 1063-5157. 72(4), p. 955–963. doi: 10.1093/sysbio/syad012. Full text in Research Archive Show summary

  Models based on the Ornstein–Uhlenbeck process have become standard for the comparative study of adaptation. Cooper et al. (2016) have cast doubt on this practice by claiming statistical problems with fitting Ornstein–Uhlenbeck models to comparative data. Specifically, they claim that statistical tests of Brownian motion may have too high Type I error rates and that such error rates are exacerbated by measurement error. In this note, we argue that these results have little relevance to the estimation of adaptation with Ornstein–Uhlenbeck models for three reasons. First, we point out that Cooper et al. (2016) did not consider the detection of distinct optima (e.g. for different environments), and therefore did not evaluate the standard test for adaptation. Second, we show that consideration of parameter estimates, and not just statistical significance, will usually lead to correct inferences about evolutionary dynamics. Third, we show that bias due to measurement error can be corrected for by standard methods. We conclude that Cooper et al. (2016) have not identified any statistical problems specific to Ornstein–Uhlenbeck models, and that their cautions against their use in comparative analyses are unfounded and misleading. [adaptation, Ornstein–Uhlenbeck model, phylogenetic comparative method.]

• Martino, Emanuela Di; Berning, Björn; Gordon, Dennis P; Kuklinski, Piotr; Liow, Lee Hsiang & Ramsfjell, Mali Hamre [Show all 12 contributors for this article] (2023). DeepBryo: A web app for AI-assisted morphometric characterization of cheilostome bryozoans. Limnology and Oceanography : Methods. doi: 10.1002/lom3.10563. Full text in Research Archive Show summary

  Abstract Bryozoans are becoming an increasingly popular study system in macroevolutionary, ecological, and paleobiological research. Members of this colonial invertebrate phylum display an exceptional degree of division of labor in the form of specialized modules, which allows for the inference of individual allocation of resources to reproduction, defense, and growth using simple morphometric tools. However, morphometric characterizations of bryozoans are notoriously labored. Here, we introduce DeepBryo, a web application for deep‐learning‐based morphometric characterization of cheilostome bryozoans. DeepBryo is capable of detecting objects belonging to six classes and outputting 14 morphological shape measurements for each object. The users can visualize the predictions, check for errors, and directly filter model outputs on the web browser. DeepBryo was trained and validated on a total of 72,412 structures in six different object classes from images of 109 different families of cheilostome bryozoans. The model shows high (> 0.8) recall and precision for zooid‐level structures. Its misclassification rate is low (~ 4%) and largely concentrated in two object classes. The model's estimated structure‐level area, height, and width measurements are statistically indistinguishable from those obtained via manual annotation. DeepBryo reduces the person‐hours required for characterizing individual colonies to less than 1% of the time required for manual annotation. Our results indicate that DeepBryo enables cost‐, labor,‐ and time‐efficient morphometric characterization of cheilostome bryozoans. DeepBryo can greatly increase the scale of macroevolutionary, ecological, taxonomic, and paleobiological analyses, as well as the accessibility of deep‐learning tools for this emerging model system.

• Voje, Kjetil Lysne (2023). Fitting and evaluating univariate and multivariate models of within-lineage evolution. Paleobiology. ISSN 0094-8373. 49(4), p. 747–764. doi: 10.1017/pab.2023.10. Full text in Research Archive Show summary

  The nature of phenotypic evolution within lineages is central to many unresolved questions in paleontology and evolutionary biology. Analyses of evolutionary time series of ancestor–descendant populations in the fossil record are likely to make important contributions to many of these debates. However, the limited number of models that have been applied to these types of data may restrict our ability to interpret phenotypic evolution in the fossil record. Using uni- and multivariate models of trait evolution that make different assumptions regarding the dynamics of the adaptive landscape, I evaluate contrasting hypotheses to explain evolution of size in the radiolarian Eucyrtidium calvertense and armor in the stickleback Gasterosteus doryssus. Body-size evolution in E. calvertense is best explained by a model in which the lineage evolves as a consequence of a shift in the adaptive landscape that coincides with the initiation of neosympatry with its sister lineage. Multivariate evolution of armor traits in a stickleback lineage (G. doryssus) shows evidence of adaptation toward independent optima on the adaptive landscape at the same time as traits change in a correlated fashion. The fitted models are available in the R package evoTS, which builds on the paleoTS framework.

• Bartoszek, Krzysztof; Fuentes-González, Jesualdo; Mitov, Venelin; Pienaar, Jason; Piwczyński, Marcin & Puchałka, Radosław [Show all 8 contributors for this article] (2023). Analytical advances alleviate model misspecification in non-Brownian multivariate comparative methods. Evolution. ISSN 0014-3820. 78(3), p. 389–400. doi: 10.1093/evolut/qpad185. Full text in Research Archive Show summary

  Abstract Adams and Collyer argue that contemporary multivariate (Gaussian) phylogenetic comparative methods are prone to favouring more complex models of evolution and sometimes rotation invariance can be an issue. Here we dissect the concept of rotation invariance and point out that, depending on the understanding, this can be an issue with any method that relies on numerical instead of analytical estimation approaches. We relate this to the ongoing discussion concerning phylogenetic principal component analysis. Contrary to what Adams and Collyer found, we do not observe a bias against the simpler Brownian motion process in simulations when we use the new, improved, likelihood evaluation algorithm employed by mvSLOUCH, which allows for studying much larger phylogenies and more complex model setups.

• Bartoszek, Krzysztof; Gonzalez, Jesualdo Fuentes; Mitov, Venelin; Pienaar, Jason; Piwczyński, Marcin & Puchalka, Radoslaw [Show all 8 contributors for this article] (2022). Model Selection Performance in Phylogenetic Comparative Methods Under Multivariate Ornstein-Uhlenbeck Models of Trait Evolution. Systematic Biology. ISSN 1063-5157. 72(2), p. 275–293. doi: 10.1093/sysbio/syac079. Full text in Research Archive Show summary

  Abstract The advent of fast computational algorithms for phylogenetic comparative methods allows for considering multiple hypotheses concerning the co-adaptation of traits and also for studying if it is possible to distinguish between such models based on contemporary species measurements. Here we demonstrate how one can perform a study with multiple competing hypotheses using mvSLOUCH by analysing two data sets, one concerning feeding styles and oral morphology in ungulates, and the other concerning fruit evolution in Ferula (Apiaceae). We also perform simulations to determine if it is possible to distinguish between various adaptive hypotheses. We find that Akaike’s information criterion corrected for small sample size has the ability to distinguish between most pairs of considered models. However, in some cases there seems to be bias towards Brownian motion or simpler Ornstein–Uhlenbeck models. We also find that measurement error and forcing the sign of the diagonal of the drift matrix for an Ornstein–Uhlenbeck process influences identifiability capabilities. It is a cliché that some models, despite being imperfect, are more useful than others. Nonetheless, having a much larger repertoire of models will surely lead to a better understanding of the natural world, as it will allow for dissecting in what ways they are wrong.

• Orr, Russell John Scott; Martino, Emanuela Di; Ramsfjell, Mali Hamre; Gordon, Dennis P.; Berning, Björn & Chowdhury, Ismael [Show all 34 contributors for this article] (2022). Paleozoic origins of cheilostome bryozoans and their parental care inferred by a new genome-skimmed phylogeny. Science Advances. 8(13). doi: 10.1126/sciadv.abm7452. Full text in Research Archive Show summary

  Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic.

• Voje, Kjetil L.; Bell, Michael A. & Stuart, Yoel E. (2022). Evolution of static allometry and constraint on evolutionary allometry in a fossil stickleback. Journal of Evolutionary Biology. ISSN 1010-061X. 35(3), p. 423–438. doi: 10.1111/jeb.13984. Full text in Research Archive Show summary

  Allometric scaling describes the relationship of trait size to body size within and among taxa. The slope of the population-level regression of trait size against body size (i.e. static allometry) is typically invariant among closely related populations and species. Such invariance is commonly interpreted to reflect a combination of developmental and selective constraints that delimit a phenotypic space into which evolution could proceed most easily. Thus, understanding how allometric relationships do eventually evolve is important to understanding phenotypic diversification. In a lineage of fossil Threespine Stickleback (Gasterosteus doryssus), we investigated the evolvability of static allometric slopes for nine traits (five armour and four non-armour) that evolved significant trait differences across 10 samples over 8500 years. The armour traits showed weak static allometric relationships and a mismatch between those slopes and observed evolution. This suggests that observed evolution in these traits was not constrained by relationships with body size, perhaps because prior, repeated adaptation to freshwater habitats by Threespine Stickleback had generated strong selection to break constraint. In contrast, for non-armour traits, we found stronger allometric relationships. Those allometric slopes did evolve on short time scales. However, those changes were small and fluctuating and the slopes remained strong predictors of the evolutionary trajectory of trait means over time (i.e. evolutionary allometry), supporting the hypothesis of allometry as constraint.

• Shaw, Josh Jenkins & Voje, Kjetil Lysne (2021). Horn scaling relationships in three species of Bledius Leach 1819 (Insecta: Coleoptera: Staphylinidae) show no indication of fitting non-linear allometric models. Journal of Natural History. ISSN 0022-2933. doi: 10.1080/00222933.2021.1891315. Full text in Research Archive Show summary

  The males of some species of the rove beetle genus Bledius Leach (1819) have a pronounced horn protruding from the anterior margin of the pronotum. The horn function is not known, however in many other horn-bearing beetles, these structures are implicated in male-male combat over females. In this study, scaling relationships are investigated for three species; B. spectabilis Kraatz, 1857, B. frisius Lohse, 1978 and B. tricornis (Herbst, 1784). Based on data collected from museum specimens, we fitted and compared different allometric models on pronotal width versus horn length in each species. A simple linear allometric model best describes the scaling relationship of horns in all three species. The horn allometry in the three Bledius species do not accordingly follow a ‘switch point’ model or a curved allometric relationship, models that are commonly reported to explain the scaling relationship of horns in many other beetle species. We therefore found no evidence of male dimorphism (minor/major phenotypes) in the species examined. The potential function of the horn in Bledius is discussed in light of similar studies.

• Love, Alan; Grabowski, Mark Walter; Houle, David; Liow, Lee Hsiang; Porto, Arthur Guimaraes Carvalho & Tsuboi, Masahito [Show all 8 contributors for this article] (2021). Evolvability in the fossil record. Paleobiology. ISSN 0094-8373. p. 1–24. doi: 10.1017/pab.2021.36. Full text in Research Archive Show summary

  Abstract The concept of evolvability—the capacity of a population to produce and maintain evolutionarily relevant variation—has become increasingly prominent in evolutionary biology. Paleontology has a long history of investigating questions of evolvability, but paleontological thinking has tended to neglect recent discussions, because many tools used in the current evolvability literature are challenging to apply to the fossil record. The fundamental difficulty is how to disentangle whether the causes of evolutionary patterns arise from variational properties of traits or lineages rather than being due to selection and ecological success. Despite these obstacles, the fossil record offers unique and growing sources of data that capture evolutionary patterns of sustained duration and significance otherwise inaccessible to evolutionary biologists. Additionally, there exist a variety of strategic possibilities for combining prominent neontological approaches to evolvability with those from paleontology. We illustrate three of these possibilities with quantitative genetics, evolutionary developmental biology, and phylogenetic models of macroevolution. In conclusion, we provide a methodological schema that focuses on the conceptualization, measurement, and testing of hypotheses to motivate and provide guidance for future empirical and theoretical studies of evolvability in the fossil record.

• Voje, Kjetil L.; Martino, Emanuela Di & Porto, Arthur Guimaraes Carvalho (2020). Revisiting a landmark study system: No evidence for a punctuated mode of evolution in Metrarabdotos. The American Naturalist. ISSN 0003-0147. 195(5), p. 899–917. doi: 10.1086/707664. Full text in Research Archive Show summary

  Is speciation generally a “special time” in morphological evolution, or are lineage-splitting events just “more of the same” where the end product happens to be two separate lineages? Data on evolutionary dynamics during anagenetic and cladogenetic events among closely related lineages within a clade are rare, but the fossil record of the bryozoan genus Metrarabdotos is considered a textbook example of a clade where speciation causes rapid evolutionary change against a backdrop of morphological stasis within lineages. Here, we point to some methodological and measurement theoretical issues in the original work on Metrarabdotos. We then reanalyze a subset of the original data that can be meaningfully investigated using quantitative statistical approaches similar to those used in the original studies. We consistently fail to find variation in the evolutionary process during within-lineage evolution compared with cladogenetic events: the rates of evolution, the strength of selection, and the directions traveled in multivariate morphospace are not different when comparing evolution within lineages and at speciation events in Metrarabdotos, and genetic drift cannot be excluded as a sufficient explanation for the morphological differentiation within lineages and during speciation. Although widely considered the best example of a punctuated mode of evolution, morphological divergence and speciation are not linked in Metrarabdotos.

• Porto, Arthur Guimaraes Carvalho & Voje, Kjetil L. (2020). ML‐morph: A fast, accurate and general approach for automated detection and landmarking of biological structures in images. Methods in Ecology and Evolution. 11(4), p. 500–512. doi: 10.1111/2041-210X.13373. Full text in Research Archive Show summary

  1. Morphometrics has become an indispensable component of the statistical analysis of size and shape variation in biological structures. Morphometric data have traditionally been gathered through low‐throughput manual landmark annotation, which represents a significant bottleneck for morphometric‐based phenomics. Here we propose a machine‐learning‐based high‐throughput pipeline to collect high‐dimensional morphometric data in two‐dimensional images of semi‐rigid biological structures. 2. The proposed framework has four main strengths. First, it allows for dense phenotyping with minimal impact on specimens. Second, it presents landmarking accuracy comparable to manual annotators, when applied to standardized datasets. Third, it performs data collection at speeds several orders of magnitude higher than manual annotators. And finally, it is of general applicability (i.e. not tied to a specific study system). 3. State‐of‐the‐art validation procedures show that the method achieves low error levels when applied to three morphometric datasets of increasing complexity, with error varying from 0.57% to 2.2% of the structure's length in the automated placement of landmarks. As a benchmark for the speed of the entire automated landmarking pipeline, our framework places 23 landmarks on 13,686 objects (zooids) detected in 1,684 pictures of fossil bryozoans in 3.12 min using a personal computer. 4. The proposed machine‐learning‐based phenotyping pipeline can greatly increase the scale, reproducibility and speed of data collection within biological research. To aid the use of the framework, we have developed a file conversion algorithm that can be used to leverage current morphometric datasets for automation, allowing the entire procedure, from model training all the way to prediction, to be performed in a matter of hours.

• Tsuboi, Masahito; Kopperud, Bjørn Tore; Syrowatka, Christine; Grabowski, Mark Walter; Voje, Kjetil L. & Pelabon, Christophe [Show all 7 contributors for this article] (2020). Measuring Complex Morphological Traits with 3D Photogrammetry: A Case Study with Deer Antlers. Evolutionary Biology. ISSN 0071-3260. 47, p. 175–186. doi: 10.1007/s11692-020-09496-9. Full text in Research Archive Show summary

  The increasing availability of 3D-imaging technology provides new opportunities for measuring morphology. Photogrammetry enables easy 3D-data acquisition compared to conventional methods and here we assess its accuracy for measuring the size of deer antlers, a complex morphological structure. Using a proprietary photogrammetry software, we generated 3D images of antlers for 92 individuals from 29 species of cervids that vary widely in antler size and shape and used these to measure antler volume. By repeating the process, we found that the relative error averaged 8.5% of object size. Errors in converting arbitrary voxel units into real volumetric units accounted for 70% of the measurement variance and can therefore be reduced by replicating the conversion. We applied the method to clay models of known volume and found no indication of bias. The estimation was robust against variation in imaging device, distance and operator, but approximately 40 images per specimen were necessary to achieve good precision. We used the method to show that conventional measures of main-beam length are relatively poor estimators of antler volume. Using loose antlers of known weight, we also showed that the volume may be a relatively poor predictor of antler weight due to variation in bone density across species. We conclude that photogrammetry can be an efficient and accurate tool for measuring antlers, and likely many other complex morphological traits.

• Liow, Lee Hsiang; Reitan, Trond; Voje, Kjetil L.; Taylor, Paul D. & Martino, Emanuela Di (2019). Size, weapons, and armor as predictors of competitive outcomes in fossil and contemporary marine communities. Ecological Monographs. ISSN 0012-9615. p. 1–15. doi: 10.1002/ecm.1354. Full text in Research Archive
• Voje, Kjetil L. (2019). Testing eco-evolutionary predictions using fossil data: Phyletic evolution following ecological opportunity. Evolution. ISSN 0014-3820. 74(1), p. 188–200. doi: 10.1111/evo.13838. Full text in Research Archive Show summary

  Fossil sequences provide observations of phenotypes within a lineage over time and represent essential data for increasing our understanding of phyletic evolution beyond microevolutionary timescales. I investigate if fossil time series of the diatom Stephanodiscus niagarae/yellowstonensis follow evolutionary dynamics compatible with hypotheses for how the adaptive landscape changes when a population enters a new environment. The lineage—which has a remarkably detailed stratigraphic record—invaded Yellowstone Lake immediately after recession of ice from the basin 14,000 years ago. Several phyletic models portraying different types of evolutionary dynamics—both compatible and not compatible with changes in the adaptive landscape following ecological opportunity—were fitted to the fossil times‐series of S. niagarae/yellowstonensis. Different models best describe the three analyzed traits. Two of the models (a new model of decelerated evolution and an Ornstein–Uhlenbeck model) capture trait dynamics compatible with an event of ecological opportunity, whereas the third model (random walk) does not. Entering a new environment may accordingly affect trait dynamics for thousands of years, but the effects can vary across phenotypes. However, tests of model adequacy reveal shortcomings in all three models explaining the trait dynamics, suggesting model development is needed to more fully understand the phyletic evolution in S. niagarae/yellowstonensis.

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• Balk, Meghan & Voje, Kjetil L. (2026). Genetic constraints unaffected by selection predicts evolution within a fossil lineage. Full text in Research Archive
• Voje, Kjetil L. (2025). Phenotypic evolution in the fossil record: Connecting microevolutionary processes to macroevolutionary patterns. Full text in Research Archive
• Voje, Kjetil L. (2025). Within-population variance predicts evolution in a coccolithophore lineage transitioning between adaptive zones on the macroevolutionary adaptive landscape. Full text in Research Archive
• Voje, Kjetil L. (2025). Phenotypic evolution in the fossil record. Full text in Research Archive
• Voje, Kjetil L. (2025). The stability of evolvability during different modes of evolution. Full text in Research Archive
• Voje, Kjetil L. (2025). Evolvability predicts evolutionary divergence across micro- and macroevolution. Full text in Research Archive
• Voje, Kjetil L. (2025). Hunndyr lever lenger enn hanndyr, men hos fuglene er det motsatt. [Internet]. forskning.no. Full text in Research Archive Show summary

  – Man kan tenke seg at alle individer har en viss mengde med energi å bruke på ulike ting, forklarer norsk forsker.

• Voje, Kjetil L. (2025). Newton: Overnatter på museum! - NRK TV. [TV]. NRK. Full text in Research Archive Show summary

  Gjør kule ting vitenskapelig og vitenskapelige ting kule

• Voje, Kjetil L. (2025). Hvorfor får vi gåsehud? – En påminnelse om at vi er dyr. [Internet]. ung.forskning.no. Full text in Research Archive Show summary

  – Hvert hår på huden har en liten muskel festet til seg, sier hudforsker.

• Voje, Kjetil L. (2025). Skoleomvisning - Naturhistorisk Museum. Full text in Research Archive
• Voje, Kjetil L. (2025). Ville jeg funnet noe å spise i perm eller jura? - Titan.uio.no. [Internet]. titan.uio.no. Full text in Research Archive Show summary

  Hvis vi kunne reist hundrevis av millioner år tilbake i tid, hva skulle vi spist for å holde oss i live?

• Voje, Kjetil L. (2025). Kan maur gå seg vill? - Titan.uio.no. [Internet]. titan.uio.no. Full text in Research Archive Show summary

  Hva skjer med en maur som havner på avveie? Finner den tilbake til tua si?

• Voje, Kjetil L. & Raklev, Are (2025). Kan KI lage en helt ny farge? - Abels tårn - NRK Radio. [Radio]. NRK. Full text in Research Archive Show summary

  Sammen med et ekspertpanel gir Torkild Jemterud svar på alt du ikke visste du lurte på Dagens eksperter: Fysiker Are Raklev Biolog Kjetil Lysne Voje Send oss dine spørsmål ved å trykke på snakkebobla øverst i NRK-radioappen, eller send mail til abels@nrk.no

• Jemterud, Torkild; Bjorå, Charlotte Sletten; Görbitz, Carl Henrik; Bøhn, Einar Duenger; Dietrichs, Erik Sveberg & Paus-Knudsen, Julie Sørlie [Show all 8 contributors for this article] (2025). Abels tårns ekstrasending. [Radio]. P2. doi: https:/radio.nrk.no/podkast/abels_taarn/sesong/2025. Full text in Research Archive
• Voje, Kjetil Lysne (2024). Evolution: Fast or Slow? Lizards Help Resolve a Paradox. [Journal]. quanta magazine. Full text in Research Archive
• Voje, Kjetil Lysne (2024). Phenotypic evolution in the fossil record. Full text in Research Archive
• Thaureau, Marion & Voje, Kjetil L. (2024). Exploring Phenotypic Evolution as a consequence of changes in the Adaptive Landscape across Lineages and Timescales. Full text in Research Archive
• Zanandrea, Irene; Voje, Kjetil L. & Balk, Meghan (2024). Assessing the importance of variational properties and developmental bias in predicting evolution in mammals. Full text in Research Archive
• Dalseng, Carmen Celine Tyndale From & Voje, Kjetil L. (2024). A phylogenetic study of Lemmus. Full text in Research Archive
• Dalseng, Carmen Celine Tyndale From & Voje, Kjetil L. (2024). A phylogenetic study of Lemmus with focus on the subspecies of Novaya Zemlya. Full text in Research Archive
• Voje, Kjetil L. (2024). Hvorfor trenger vi et artsmangfold, og hva finnes av artsmangfold her i byen? Full text in Research Archive
• Voje, Kjetil L. (2024). Samlingene - blikk inn i naturens gullgruve. Full text in Research Archive
• Voje, Kjetil L. (2024). Hvorfor trenger vi et artsmangfold, og hva finnes av artsmangfold her i byen? Full text in Research Archive
• Voje, Kjetil L. (2024). Hvorfor trenger vi et artsmangfold, og hva finnes av artsmangfold her i byen? Full text in Research Archive
• Balk, Meghan & Voje, Kjetil L. (2024). Is phenotypic evolution predicted by directions of high trait variation during anagenesis and cladogenesis? Full text in Research Archive
• Balk, Meghan & Voje, Kjetil L. (2024). The role of trait variation in predicting directions phenotypic evolution during anagenesis and cladogenesis. Full text in Research Archive
• Thaureau, Marion & Voje, Kjetil L. (2024). EXPLORING PHENOTYPIC EVOLUTION AS A CONSEQUENCE OF CHANGES IN THE ADAPTIVE LANDSCAPE ACROSS LINEAGES AND TIMESCALES. Full text in Research Archive
• Voje, Kjetil L. (2024). Hvorfor har vi samlinger? Full text in Research Archive
• Thaureau, Marion; Kinneberg, Vilde Bruhn & Voje, Kjetil L. (2024). Beyond the traditional three: Assessing relative model fit of several evolutionary models to fossil time series. Full text in Research Archive
• Thaureau, Marion & Voje, Kjetil L. (2024). Phenotypic Evolution as a consequence of changes in the Adaptive Landscape across Lineages and Timescales. Full text in Research Archive
• Thaureau, Marion; Rugstad, Audun & Voje, Kjetil L. (2024). The Phenotypic Evolution Time Series (Pets) database: facilitating research on phenotypic change within lineages. Full text in Research Archive
• Lifjeld, Jan Terje; Cramer, Becky; Leder, Erica Helen & Voje, Kjetil L. (2024). Sperm as a speciation phenotype in songbirds. Full text in Research Archive
• Thaureau, Marion & Voje, Kjetil L. (2023). Covariation of phenotypic traits and environmental factors across timescales and lineages. Full text in Research Archive
• Voje, Kjetil L. (2023). Voje samler rådata fra disketter og harddisker. [Journal]. titan.uio.no. Full text in Research Archive
• Voje, Kjetil L. (2023). Disse to hundene har omtrent like stor hjerne, men hva har størrelse å si for intelligensen? [Journal]. forskning.no. Full text in Research Archive
• Voje, Kjetil L. (2023). Arter som var vanlige før finnes bare på museum i dag. [Internet]. titan.uio.no. Full text in Research Archive
• Voje, Kjetil L. (2023). A NEW MODELING FRAMEWORK FOR FITTING AND EVALUATING UNIVARIATE AND MULTIVARIATE MODELS OF PHYLETIC EVOLUTION. Full text in Research Archive
• Voje, Kjetil L. (2023). Evolvability in the fossil record. Full text in Research Archive
• Voje, Kjetil L. (2023). EVOLVABILITY PREDICTS EVOLUTIONARY DIVERGENCE ACROSS MICRO- AND MACROEVOLUTION. Full text in Research Archive
• Voje, Kjetil L. (2023). Document your data, share your data. Full text in Research Archive
• Voje, Kjetil L. (2023). Leter i skuffer og skap. [Journal]. Teknisk Ukeblad. Full text in Research Archive
• Thaureau, Marion & Voje, Kjetil L. (2023). EXPLORING PHENOTYPIC EVOLUTION AS A CONSEQUENCE OF CHANGES IN THE ADAPTIVE LANDSCAPE ACROSS LINEAGES AND TIMESCALES. Full text in Research Archive
• Voje, Kjetil L. (2023). Den evolusjonære svarte boksen. [Journal]. Biolog. Full text in Research Archive
• Voje, Kjetil L. (2023). Søkelys på truet natur. [Journal]. Palmebladet. Full text in Research Archive
• Voje, Kjetil L. (2023). Wallace, Darwin og naturlig seleksjon. Full text in Research Archive
• Voje, Kjetil L. (2023). Evolvability in the fossil record - A case study. Full text in Research Archive
• Voje, Kjetil L. (2023). Hva er evolusjonsbiologi? Full text in Research Archive
• Balk, Meghan & Voje, Kjetil Lysne (2023). Assessing evolvability in a fossil lineage using a CV pipeline to produce a phenomic dataset. Full text in Research Archive
• Voje, Kjetil L. (2023). Abels Tårn - 29. september. [Radio]. NRK. Full text in Research Archive
• Voje, Kjetil L. (2023). Abels Tårn. [Radio]. NRK. Full text in Research Archive
• Voje, Kjetil L. (2023). Abels Tårn - 7. april 2023. [Radio]. NRK. Full text in Research Archive
• Rugstad, Audun & Voje, Kjetil L. (2023). The Phenotypic Evolution Time Series (PETS) Database: Facilitating Research on Phenotypic Change Within Lineages. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Evolusjonsbiologi. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Evolusjonsbiologi. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Inferring the dynamics of the adaptive landscape using fossil time-series. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Kan rovdyr spise planter hvis de ikke har tilgang på annen mat? [Journal]. Titan.uio.no. Full text in Research Archive
• Voje, Kjetil Lysne (2022). A NEW MODELING FRAMEWORK FOR FITTING AND EVALUATING UNIVARIATE AND MULTIVARIATE MODELS OF PHYLETIC EVOLUTION. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Evolution and evolvability in the fossil record. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Measurement theory and paleobiology. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Hvorfor legger høna egg året rundt? [Journal]. Titan.uio.no. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Evolusjon på mikronivå - gener. Full text in Research Archive
• Voje, Kjetil Lysne (2022). A number is not a number:  The importance of measurement theory. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Slik er ulvenes liv etter døden. [Journal]. Dagsavisen. Full text in Research Archive
• Voje, Kjetil Lysne (2022). Abels tårn. [Journal]. NRK.no. Full text in Research Archive Show summary

  https://radio.nrk.no/serie/abels-taarn-radio/sesong/202204/MDFP05002722

• Martino, Emanuela Di; Berning, Björn; Gordon, Dennis P.; Kuklinski, Piotr; Liow, Lee Hsiang & Ramsfjell, Mali Hamre [Show all 11 contributors for this article] (2022). DeepBryo: a web app for AI-assisted morphometric characterization of cheilostome bryozoan colonies. Full text in Research Archive
• Voje, Kjetil L. & Sofie, Frøyland Ida (2021). Noen dyr kan få unger selv om kroppen ikke har blitt voksen. [Internet]. forskning.no. Full text in Research Archive
• Voje, Kjetil L. & Vogt, Yngve (2021). Forskeren løser evolusjonsgåte med kunstig intelligens. [Journal]. Teknisk ukeblad. Full text in Research Archive
• Torkild, Jemterud & Voje, Kjetil L. (2021). Går IQ i arv? [Radio]. NRK. Full text in Research Archive
• Voje, Kjetil L. & Torgersen, Eivind (2021). Hvorfor lager elgen ovale kuler når den bæsjer? [Internet]. Titan. Full text in Research Archive
• Voje, Kjetil L. & Vogt, Yngve (2021). Forskere bruker kunstig intelligens for å forstå mer om rask og sakte evolusjon. [Internet]. Titan. Full text in Research Archive
• Torgersen, Eivind & Voje, Kjetil L. (2021). Hvorfor skriker en baby så forferdelig høyt og skjærende? [Internet]. Titan. Full text in Research Archive
• Tsuboi, Masahito; Bijl, Wouter van der; Kopperud, Bjørn Tore; Erritzøe, Johannes; Voje, Kjetil L. & Kotrschal, Alexander [Show all 10 contributors for this article] (2019). Reply to: Comparisons of static brain–body allometries across vertebrates must distinguish between indeterminate and determinate growth. Nature Ecology and Evolution. 3(10), p. 1405–1406. doi: 10.1038/s41559-019-0986-z. Full text in Research Archive

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