Evolutionary Transitions Research Articles

Ultra-high Q resonances based on zero group-velocity modes accompanied by bound states in the continuum in 2D photonic crystal slabs

Optical resonators made of 2D photonic crystal (PhC) slabs provide efficient ways to manipulate light at the nanoscale through small group-velocity modes with low radiation losses. The resonant modes in periodic photonic lattices are predominantly limited by nonleaky guided modes at the boundary of the Brillouin zone below the light cone. Here, we propose a mechanism for ultra-high Q resonators based on the bound states in the continuum (BICs) above the light cone that have zero-group velocity (ZGV) at an arbitrary Bloch wavevector. By means of the mode expansion method, the construction and evolution of avoided crossings and Friedrich-Wintgen BICs are theoretically investigated at the same time. By tuning geometric parameters of the PhC slab, the coalescence of eigenfrequencies for a pair of BIC and ZGV modes is achieved, indicating that the waveguide modes are confined longitudinally by small group-velocity propagation and transversely by BICs. Using this mechanism, we engineer ultra-high Q nanoscale resonators that can significantly suppress the radiative losses, despite the operating frequencies above the light cone and the momenta at the generic k point. Our work suggests that the designed devices possess potential applications in low-threshold lasers and enhanced nonlinear effects.

Trade-Offs in the Sensory Brain between Diurnal and Nocturnal Rodents

Introduction: Transitions in temporal niche have occurred many times over the course of mammalian evolution. These are associated with changes in sensory stimuli available to animals, particularly with visual cues, because levels of light are so much higher during the day than at night. This relationship between temporal niche and available sensory stimuli elicits the expectation that evolutionary transitions between diurnal and nocturnal lifestyles will be accompanied by modifications of sensory systems that optimize the ability of animals to receive, process, and react to important stimuli in the environment. Methods: This study examines the influence of temporal niche on investment in sensory brain tissue of 13 rodent species (five diurnal; eight nocturnal). Animals were euthanized and the brains immediately frozen on dry ice; olfactory bulbs were subsequently dissected and weighed, and the remaining brain was weighed, sectioned, and stained. Stereo Investigator was used to calculate volumes of four sensory regions that function in processing visual (lateral geniculate nucleus, superior colliculus) and auditory (medial geniculate nucleus, inferior colliculus) information. A phylogenetic framework was used to assess the influence of temporal niche on the relative sizes of these brain structures and of olfactory bulb weights. Results: Compared to nocturnal species, diurnal species had larger visual regions, whereas nocturnal species had larger olfactory bulbs than their diurnal counterparts. Of the two auditory structures examined, one (medial geniculate nucleus) was larger in diurnal species, while the other (inferior colliculus) did not differ significantly with temporal niche. Conclusion: Our results indicate a possible indirect association between temporal niche and auditory investment and suggest probable trade-offs of investment between olfactory and visual areas of the brain, with diurnal species investing more in processing visual information and nocturnal species investing more in processing olfactory information.

Natural variation in the Caenorhabditis elegans egg-laying circuit modulates an intergenerational fitness trade-off.

Evolutionary transitions from egg laying (oviparity) to live birth (viviparity) are common across various taxa. Many species also exhibit genetic variation in egg-laying mode or display an intermediate mode with laid eggs containing embryos at various stages of development. Understanding the mechanistic basis and fitness consequences of such variation remains experimentally challenging. Here, we report highly variable intra-uterine egg retention across 316 Caenorhabditis elegans wild strains, some exhibiting strong retention, followed by internal hatching. We identify multiple evolutionary origins of such phenotypic extremes and pinpoint underlying candidate loci. Behavioral analysis and genetic manipulation indicates that this variation arises from genetic differences in the neuromodulatory architecture of the egg-laying circuitry. We provide experimental evidence that while strong egg retention can decrease maternal fitness due to in utero hatching, it may enhance offspring protection and confer a competitive advantage. Therefore, natural variation in C. elegans egg-laying behaviour can alter an apparent trade-off between different fitness components across generations. Our findings highlight underappreciated diversity in C. elegans egg-laying behavior and shed light on its fitness consequences. This behavioral variation offers a promising model to elucidate the molecular changes in a simple neural circuit underlying evolutionary shifts between alternative egg-laying modes in invertebrates.

Natural variation in the Caenorhabditis elegans egg-laying circuit modulates an intergenerational fitness trade-off

Evolutionary transitions from egg laying (oviparity) to live birth (viviparity) are common across various taxa. Many species also exhibit genetic variation in egg-laying mode or display an intermediate mode with laid eggs containing embryos at various stages of development. Understanding the mechanistic basis and fitness consequences of such variation remains experimentally challenging. Here, we report highly variable intra-uterine egg retention across 316 Caenorhabditis elegans wild strains, some exhibiting strong retention, followed by internal hatching. We identify multiple evolutionary origins of such phenotypic extremes and pinpoint underlying candidate loci. Behavioral analysis and genetic manipulation indicates that this variation arises from genetic differences in the neuromodulatory architecture of the egg-laying circuitry. We provide experimental evidence that while strong egg retention can decrease maternal fitness due to in utero hatching, it may enhance offspring protection and confer a competitive advantage. Therefore, natural variation in C. elegans egg-laying behaviour can alter an apparent trade-off between different fitness components across generations. Our findings highlight underappreciated diversity in C. elegans egg-laying behavior and shed light on its fitness consequences. This behavioral variation offers a promising model to elucidate the molecular changes in a simple neural circuit underlying evolutionary shifts between alternative egg-laying modes in invertebrates.

The neoteny goldilocks zone: The evolution of neoteny in Ambystoma.

Neoteny is a developmental strategy wherein an organism reaches sexual maturity without associated adult characteristics. In salamanders, neoteny takes the form of individuals retaining aquatic larval characteristics such as external gills upon maturation. Mole salamanders (Ambystoma) occupy a wide range of habitats and areas across the North American continent, and display examples of non-neotenic, facultatively neotenic and obligate neotenic species, providing high variation for investigating the factors influencing the evolution of neoteny. Here, we use phylogenetic comparative methods to test existing hypotheses that neoteny is associated with elevational and latitudinal distribution, cave-associated isolation, and hybridisation-related polyploidy. We also test if neoteny influences the diversity of habitats a species can occupy, since the restriction to an aquatic life should constrain the availability of different niches. We find that neoteny tends to occur in a narrow latitudinal band between 20-30° North, with particularly narrow latitudinal ranges for obligate compared to facultative neotenic species (16-52° North). We also find that facultatively neotenic species occur at elevations more than twice as high as other species on average, and that species with a higher frequency of neoteny typically have lower habitat diversity. Our results suggest that evolutionary transitions between non-neotenic and facultative neoteny states occur at relatively high and approximately equal rates. Moreover, we estimate that obligate neoteny cannot evolve directly from non-neotenic species (and vice versa), such that facultative neoteny acts as an evolutionary 'stepping stone' to and from obligate neoteny. However, our transition rate estimates suggest that obligate neoteny is lost >4-times faster than it evolves, partly explaining the rarity of obligate species. These results support the hypothesis that low latitudes favour the evolution of neoteny, presumably linked to more stable (aquatic) environments due to reduced seasonality, but once evolved it may constrain the diversity of habitats.

Anharmonic effects on the squeezing of axion perturbations

It is assumed in standard cosmology that the Universe underwent a period of inflation in its earliest phase, providing the seeds for structure formation through vacuum fluctuations of the inflaton scalar field. These fluctuations get stretched by the quasi-exponential expansion of the Universe and become squeezed. The aim of this paper is to deepen the understanding of the squeezing process, considering the effect of self-interactions. Axion-like particles can provide a useful setup to study this effect. Specifically we focus on the consequences that a non-trivial evolution of the background axion field has on the squeezing of the perturbations. We follow the evolution of the axion's fluctuation modes from the horizon exit during inflation to the radiation-dominated epoch. We compute Bogoliubov coefficients and squeezing parameters, which are linked to the axion particle number and isocurvature perturbation. We find that the quantum mechanical particle production and the squeezing of the perturbations are enhanced, if one accounts for anharmonic effects, i.e., the effect of higher order terms in the potential. This effect becomes particularly strong towards the hilltop of the potential.

Scheduling of Continuous Annealing With a Multi-Objective Differential Evolution Algorithm Based on Deep Reinforcement Learning

This paper studies a multi-objective scheduling problem in a continuous annealing operation of the steel industry, which is to simultaneously determine the production batch sizes for coils and their schedules so as to minimize the cost of setup, earliness, and tardiness. A large batch can reduce the setup costs but lead to a cost increase in earliness and tardiness. Thus, the conflict objectives of minimizing setup cost, earliness, and tardiness can be formulated separately. In this paper, we formulate a multi-objective optimization model for the problem and develop an adaptive multi-objective differential evolutionary based on deep reinforcement learning (AMODE-DRL) for effectively obtaining the Pareto solutions. In the proposed AMODE-RDL, DRL is integrated into the MODE algorithm as a controller, which can adaptively select mutation operators and parameters according to different search domains. Computational results on randomly generated instances and the practical problem instances show that DRL can effectively guide MODE to select mutation operators and parameters. The proposed algorithm can obtain better solutions compared to other powerful multi-objective evolutionary and adaptive MODE algorithms. <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">Note to Practitioners</i> —Setup usually leads to a decrease in production capacity and an increase in production costs. Similarly, earliness and tardiness costs are also crucial. These costs correspond to production, customer demand, and inventory, which are common objectives in production systems. However, the three objectives are generally conflicting, and it is very hard for practitioners to make appropriate decisions with manual experience. The multi-objective optimization methods can provide different scheduling for practitioners who may choose the suitable one according to current working conditions. Accordingly, the proposed model and algorithm can extend to other fields with similar characteristic problems.

Origin and evolution of a new tetraploid mangrove species in an intertidal zone

Polyploidy is a major factor in the evolution of plants, yet we know little about the origin and evolution of polyploidy in intertidal species. This study aimed to identify the evolutionary transitions in three true-mangrove species of the genus Acanthus distributed in the Indo–West Pacific region. For this purpose, we took an integrative approach that combined data on morphology, cytology, climatic niche, phylogeny, and biogeography of 493 samples from 42 geographic sites. Our results show that the Acanthus ilicifolius lineage distributed east of the Thai–Malay Peninsula possesses a tetraploid karyotype, which is morphologically distinct from that of the lineage on the west side. The haplotype networks and phylogenetic trees for the chloroplast genome and eight nuclear genes reveal that the tetraploid species has two sub-genomes, one each from A. ilicifolius and A.ebracteatus, the paternal and maternal parents, respectively. Population structure analysis also supports the hybrid speciation history of the new tetraploid species. The two sub-genomes of the tetraploid species diverged from their diploid progenitors during the Pleistocene. Environmental niche models revealed that the tetraploid species not only occupied the near-entire niche space of the diploids, but also expanded into novel environments. Our findings suggest that A. ilicifolius species distributed on the east side of the Thai–Malay Peninsula should be regarded as a new species, A. tetraploideus, which originated from hybridization between A. ilicifolius and A. ebracteatus, followed by chromosome doubling. This is the first report of a true-mangrove allopolyploid species that can reproduce sexually and clonally reproduction, which explains the long-term adaptive potential of the species.

Investigating Small Disturbance Stability in Wind Farm Grid-Connected Systems: A Data-Driven Approach

The presence of unknown synchronization characteristics, unclear instability mechanism, and various fault mode evolution laws, lacking corresponding theoretical support and analysis methods and instability criteria, are defined with clear physical concepts. It is still impossible to systematically understand the transient synchronization mechanism of the wind power grid-connected system from the perspective of the whole fault stage. Therefore, this study uniformly reveals its temporary synchronous stability problem and proposes a large/small disturbance adaptive synchronous stability control method, which improves the dynamic characteristics of the wind turbine through the control of the inverter itself to improve the system stability—using different scenarios, such as single doubly-fed wind turbines. The experimental results show that the small disturbance on the AC side significantly impacts the system characteristics, followed by a bit of annoyance on the DC side. The DC side fault will cause a change in system frequency characteristics, especially at the receiving end. However, compared with the Voltage Source Converters-High Voltage Direct Current (VSC-HVDC)system, Modular Multilevel Converters-High Voltage Direct Current (MMC-HVDC) systems operate at a much higher frequency and produce less low-frequency harmonics. This makes them less likely to induce subsynchronous oscillations in the system.

Plasticity-led and mutation-led evolutions are different modes of the same developmental gene regulatory network.

The standard theory of evolution proposes that mutations cause heritable variations, which are naturally selected, leading to evolution. However, this mutation-led evolution (MLE) is being questioned by an alternative theory called plasticity-led evolution (PLE). PLE suggests that an environmental change induces adaptive phenotypes, which are later genetically accommodated. According to PLE, developmental systems should be able to respond to environmental changes adaptively. However, developmental systems are known to be robust against environmental and mutational perturbations. Thus, we expect a transition from a robust state to a plastic one. To test this hypothesis, we constructed a gene regulatory network (GRN) model that integrates developmental processes, hierarchical regulation, and environmental cues. We then simulated its evolution over different magnitudes of environmental changes. Our findings indicate that this GRN model exhibits PLE under large environmental changes and MLE under small environmental changes. Furthermore, we observed that the GRN model is susceptible to environmental or genetic fluctuations under large environmental changes but is robust under small environmental changes. This indicates a breakdown of robustness due to large environmental changes. Before the breakdown of robustness, the distribution of phenotypes is biased and aligned to the environmental changes, which would facilitate rapid adaptation should a large environmental change occur. These observations suggest that the evolutionary transition from mutation-led to plasticity-led evolution is due to a developmental transition from robust to susceptible regimes over increasing magnitudes of environmental change. Thus, the GRN model can reconcile these conflicting theories of evolution.

Macroevolutionary dynamics of gene family gain and loss along multicellular eukaryotic lineages

The gain and loss of genes fluctuate over evolutionary time in major eukaryotic clades. However, the full profile of these macroevolutionary trajectories is still missing. To give a more inclusive view on the changes in genome complexity across the tree of life, here we recovered the evolutionary dynamics of gene family gain and loss ranging from the ancestor of cellular organisms to 352 eukaryotic species. We show that in all considered lineages the gene family content follows a common evolutionary pattern, where the number of gene families reaches the highest value at a major evolutionary and ecological transition, and then gradually decreases towards extant organisms. This supports theoretical predictions and suggests that the genome complexity is often decoupled from commonly perceived organismal complexity. We conclude that simplification by gene family loss is a dominant force in Phanerozoic genomes of various lineages, probably underpinned by intense ecological specializations and functional outsourcing.

Palaeoatmosphere facilitates a gliding transition to powered flight in the Eocene bat, Onychonycteris finneyi

The evolutionary transition to powered flight remains controversial in bats, the only flying mammals. We applied aerodynamic modeling to reconstruct flight in the oldest complete fossil bat, the archaic Onychonycteris finneyi from the early Eocene of North America. Results indicate that Onychonycteris was capable of both gliding and powered flight either in a standard normodense aerial medium or in the hyperdense atmosphere that we estimate for the Eocene from two independent palaeogeochemical proxies. Aerodynamic continuity across a morphological gradient is further demonstrated by modeled intermediate forms with increasing aspect ratio (AR) produced by digital elongation based on chiropteran developmental data. Here a gliding performance gradient emerged of decreasing sink rate with increasing AR that eventually allowed applying available muscle power to achieve level flight using flapping, which is greatly facilitated in hyperdense air. This gradient strongly supports a gliding (trees-down) transition to powered flight in bats.

Genome-Wide Analysis of the GLK Gene Family and Its Expression at Different Leaf Ages in the Citrus Cultivar Kanpei.

The GLK gene family plays a crucial role in the regulation of chloroplast development and participates in chlorophyll synthesis. However, the precise mechanism by which GLK contributes to citrus's chlorophyll synthesis remains elusive. The GLK gene family causes variations in the photosynthetic capacity and chlorophyll synthesis of different citrus varieties. In this study, we identified tissue-specific members and the key CcGLKs involved in chlorophyll synthesis. A total of thirty CcGLK transcription factors (TFs) were discovered in the citrus genome, distributed across all nine chromosomes. The low occurrence of gene tandem duplication events and intronic variability suggests that intronic variation may be the primary mode of evolution for CcGLK TFs. Tissue-specific expression patterns were observed for various GLK family members; for instance, CcGLK12 and CcGLK15 were specifically expressed in the skin, while CcGLK30 was specific to the ovary, and CcGLK10, CcGLK6, CcGLK21, CcGLK2, CcGLK18, CcGLK9, CcGLK28, and CcGLK8 were specifically expressed in the leaves. CcGLK4, CcGLK5, CcGLK11, CcGLK23, CcGLKl7, CcGLK26, and CcGLK20 may participate in the regulation of the ALA, prochlorophylate, protoporphyrin IX, Mg-protoporphyrin IX, Chl b, T-Chl, MG-ProtoIX ME, and POR contents in citrus.

Primitive purine biosynthesis connects ancient geochemistry to modern metabolism.

An unresolved question in the origin and evolution of life is whether a continuous path from geochemical precursors to the majority of molecules in the biosphere can be reconstructed from modern-day biochemistry. Here we identified a feasible path by simulating the evolution of biosphere-scale metabolism, using only known biochemical reactions and models of primitive coenzymes. We find that purine synthesis constitutes a bottleneck for metabolic expansion, which can be alleviated by non-autocatalytic phosphoryl coupling agents. Early phases of the expansion are enriched with enzymes that are metal dependent and structurally symmetric, supporting models of early biochemical evolution. This expansion trajectory suggests distinct hypotheses regarding the tempo, mode and timing of metabolic pathway evolution, including a late appearance of methane metabolisms and oxygenic photosynthesis consistent with the geochemical record. The concordance between biological and geological analyses suggests that this trajectory provides a plausible evolutionary history for the vast majority of core biochemistry.

Zygotic quiescence prolongs the reproductive cycle in Berchemia sinica (Rhamnaceae)

More than two-thirds of the species in Berchemia exhibit remarkable reproductive phenology, characterized by prolonged fruit ripening and the occurrence of overlapping fruit ripening and flowering. However, the underlying reasons for these phenomena remain unknown. Therefore, this study employed routine paraffin section technology, histochemical technique and scanning electron microscopy to investigate embryo sac development, differences between male and hermaphroditic flowers, as well as the overwintering strategy in Berchemia sinica. The findings of this study revealed a significant reduction in pollen viability and the number of pollen grains per flower in hermaphroditic compared to male flowers. Following fertilization, the ovary of hermaphroditic flowers does not undergo significant enlargement but gradually enters the state of zygotic quiescence. Zygotic quiescence prolongation is the main reason for the long reproductive cycle of B. sinica, and it takes approximately 14 months for the entire cycle from flower bud differentiation to fruit ripening to complete. Long reproductive cycle and complex overwintering mechanism exhibited by B. sinica resemble those seen in species belonging to Betulaceae, Fagaceae, Tapisciaceae, and Theaceae, suggesting convergent evolution during Earth's glacial period resulted in similar adaptive structures among these groups delayed fruit ripening leads to overlap between current year's flowers with previous year's fruits. The trade-off between flower and fruit for reproductive resources may have driven evolutionary transition from ancestral hermaphroditism towards androdioecy in Berchemia.

A stochastic Fokker–Planck–Master model for diatomic rarefied gas flows

The direct simulation Monte Carlo (DSMC) method is widely used for numerical solutions of the Boltzmann equation. However, the associated computational cost becomes prohibitive in the near-continuum regime. To address this limitation, the particle-based Fokker–Planck (FP) method has been extensively studied in the past decade. The FP equation, which describes Brownian motion, does not require resolution of the collisional time and length scales. While several monatomic FP models have been proposed, the modeling of diatomic gases within the FP framework has received limited attention. In this paper, we propose a new diatomic kinetic model, named the Fokker–Planck–Master (FPM) model, which can accurately describe energy exchanges between translational-rotational and translational-vibrational modes. The FPM model combines the FP equation to describe the evolution of translational and rotational modes, and the master equation to describe the evolution of the vibrational modes. The numerical test cases include relaxation problems, Couette flows, and hypersonic flows past a vertical flat plate. The results demonstrate that the FPM model shows good agreement with both analytical and DSMC solutions.

Nondispersive evolution of the pump mode in down-conversion processes and in the presence of Kerr nonlinearity

Nondispersive evolution of the pump mode in down-conversion processes and in the presence of Kerr nonlinearity

Pioneering historical LCA - A perspective on the development of personal carbon footprint 1860–2020 in Denmark

Mitigating climate change by reducing anthropogenic greenhouse gas emissions is considered an urgent challenge for society. In this context, different philosophies exist. A classic debate, referred to as “boomsters vs. doomsters”, asks whether humanity's drive for technological innovations will mitigate climate change, or whether it is justifiable to compromise our living standards, affluence, and consumption patterns. A Danish research project has elucidated this topic through cross-collaboration between historians and engineers by quantifying the personal consumption-based carbon footprint (CF) of Danish citizens, taking into account the consumption segments of food, transport, housing, and furniture. Quantification is conducted through life-cycle assessments (LCA) of five decadal timesteps during the period 1860–2020. As only one other study conducting historical (retrospective) LCA with an extensive time horizon was identified in the scientific literature, our own study provides novel insights by exploring possible approaches in this field. Historical LCA has many similarities with prospective (future) LCA, providing inspiration to the approach. The main challenge of the study is to adjust both foreground and background systems in the inventory modelling phase. Data is largely based on national statistics, books, museum case studies, expert testimonials, and proxies such as using the steam turbine as an indicator of improvements to technological efficiency over time. The results show a slightly declining carbon footprint in the period 1860–1930, from 7.7 to 6.6 t CO2-eq per capita annually. Unsurprisingly the footprint peak is not found until the 1970s, at 11 t, reducing to 5.4 t in the 2010s. The residential energy consumption segment contributes half of this CF, followed by transportation by road and rail. Discussion of the results shows that whether technology can mitigate the impact from the increase in affluence depends on the specific consumption segment. For housing combined with residential energy, the results show that technology can mitigate increases in CF, despite the comforts of home having only increased over time. For the transportation segment, however, the past century has seen an eightfold increase in CF due to the pronounced evolution of personal modes of transportation. The findings suggests that the “boomsters vs. doomsters” debate is not so black and white after all. We further suggest that historical LCA, in tandem with prospective LCA, can provide valuable insights for decision support to facilitate a green transition and hence a green future.

Dominance between self-incompatibility alleles determines the mating system of Capsella allopolyploids.

The shift from outcrossing to self-fertilization is one of the main evolutionary transitions in plants and has broad effects on evolutionary trajectories. In Brassicaceae, the ability to inhibit self-fertilization is controlled by 2 genes, SCR and SRK, tightly linked within the S-locus. A series of small non-coding RNAs also encoded within the S-locus regulates the transcriptional activity of SCR alleles, resulting in a linear dominance hierarchy between them. In Brassicaceae, natural allopolyploid species are often self-compatible (SC) even when one of the progenitor species is self-incompatible, but the reason why polyploid lineages tend to lose self-incompatibility (SI) and the timing of the loss of SI (immediately after ancestral hybridization between the progenitor species, or at a later stage after the formation of allopolyploid lineages) have generally remained elusive. We used a series of synthetic diploid and tetraploid hybrids obtained between self-fertilizing Capsella orientalis and outcrossing Capsella grandiflora to test whether the breakdown of SI could be observed immediately after hybridization, and whether the occurrence of SC phenotypes could be explained by the dominance interactions between S-haplotypes inherited from the parental lineages. We used RNA-sequencing data from young inflorescences to measure allele-specific expression of the SCR gene and infer dominance interactions in the synthetic hybrids. We then evaluated the seed set from autonomous self-pollination in the synthetic hybrids. Our results demonstrate that self-compatibility of the hybrids depends on the relative dominance between S-alleles inherited from the parental species, confirming that SI can be lost instantaneously upon formation of the ancestral allopolyploid lineage. They also confirm that the epigenetic regulation that controls dominance interactions between S-alleles can function between subgenomes in allopolyploids. Together, our results illustrate how a detailed knowledge of the mechanisms controlling SI can illuminate our understanding of the patterns of co-variation between the mating system and changes in ploidy.

A new sphenodontian (Diapsida: Lepidosauria) from the Upper Triassic (Norian) of Germany and its implications for the mode of sphenodontian evolution

The Arnstadt Formation of Saxony-Anhalt, Germany has yielded some of Germany’s most substantial finds of Late Triassic tetrapods, including the sauropodomorph Plateosaurus and the stem-turtle Proganochelys quenstedti. Here, we describe an almost complete skull of a new sphenodontian taxon from this formation (Norian, 227–208 Ma), making it the oldest known articulated sphenodontian skull from Europe and one of the oldest in the world. The material is represented by the dermal skull roof and by the complete maxilla and temporal region, as well as parts of the palate, braincase, and lower jaw. A phylogenetic assessment recovers it as a basal sphenodontian closely related to Planocephalosaurus robinsonae and to Eusphenodontia, making it the earliest-diverging sphenodontian known with an articulated skull. Its cranial anatomy is generally similar to the well-known Diphydontosaurus avonis from the Rhaetian of England, showing that this successful phenotype was already established in the clade around 10 myr earlier than assumed. An analysis of evolutionary change rates recovers high rates of evolution in basal sphenodontians, with decreasing rates throughout the evolution of the group. However, contrary to previous studies, reversals in this trend were identified, indicating additional peaks of evolutionary change. These results improve our understanding of the early sphenodontian diversity in Europe, providing critical information on evolutionary trends throughout the history of the clade and sparking renewed interest in its evolution.