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Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov ., Clambiformia ser. nov. and Clamboidea sensu nov. , Rhinorhipiformia ser. nov ., Byrrhoidea sensu nov. , Dryopoidea stat. res. , Nosodendriformia ser. nov. and Staphyliniformia sensu nov ., and Erotyloidea stat. nov ., Nitiduloidea stat. nov . and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic–Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.

High-throughput sequencing projects generate genome-scale sequence data for species level phylogenies. However, state-of-the-art Bayesian methods for inferring timetrees are computationally limited to small datasets and cannot exploit the deluge of new genomes. In the case of mammals, molecular-clock analyses of limited datasets have produced conflicting estimates of clade ages with large uncertainties and, thus, the timescale of placental mammal evolution remains contentious. Here we develop a Bayesian molecular-clock dating approach to estimate a timetree of 4,705 mammal species integrating information from 72 mammal genomes. We show increasingly larger phylogenomic datasets produce diversification time estimates with progressively smaller uncertainties, facilitating precise tests of macroevolutionary hypotheses. For example, we confidently reject an explosive model of placental mammal origination in the Paleogene8 and show crown Placentalia originated in the Late Cretaceous with unambiguous ordinal diversification in the Paleocene/Eocene. Our new Bayesian methodology facilitates analysis of complete genomes and thousands of species within an integrated framework, making it possible to address hitherto intractable research questions on species diversifications. As such, our approach can be used to tackle other contentious cases of animal and plant diversifications that require analysis of species-level phylogenomic datasets.

Molecular timescales estimate that early animal lineages diverged tens of millions of years before their earliest unequivocal fossil evidence. The Ediacaran macrobiota (~574–538 million years ago) are largely eschewed from this debate, primarily due to their extreme phylogenetic uncertainty, but they remain germane. We characterize the development of Charnia masoni and establish the phylogenetic affinity of rangeomorphs, among the oldest and most enigmatic components of the Ediacaran macrobiota. We provide the first direct evidence for the internal interconnected nature of rangeomorphs, and show Charnia was constructed of repeated branches that derived successively from pre-existing branches. We find homology and rationalise morphogenesis between disparate rangeomorph taxa, before producing a new phylogenetic analysis, resolving Charnia as a stem-eumetazoan and expanding the anatomical disparity of that group to include a long-extinct bodyplan. These data bring competing records of early animal evolution into closer agreement, reformulating our understanding of the evolutionary emergence of animal bodyplans.

Fleas (Siphonaptera) are medically important blood-feeding insects responsible for spreading pathogens such as plague, murine typhus, and myxomatosis. The peculiar morphology of fleas resulting from their specialised ectoparasitic lifestyle has meant that the phylogenetic position of this diverse and medically important group has remained one of the most persistent problems in insect evolution. Here we test competing hypotheses on the contentious evolutionary relationships of fleas and antliophoran insects using the largest molecular dataset available to date consisting of over 1,400 protein-coding genes, and a smaller mitogenome and Sanger sequence alignment of 16 genes. By removing ambiguously aligned sequence regions and using site-heterogeneous models, we consistently recover fleas nested within scorpionflies (Mecoptera) as sister to the relictual southern hemisphere family Nannochoristidae. Topology tests accounting for compositional heterogeneity strongly favour the proposed topology over previous hypotheses of antliophoran relationships. This clade is diagnosed by shared morphological characters of the head and sperm pump. Fleas may no longer be regarded as a separate insect order and we propose that Siphonaptera should be treated as an infraorder within Mecoptera, reducing the number of extant holometabolan insect orders to ten.

Western honey bees (Apis mellifera) are one of the most important pollinators of agricultural crops and wild plants. Despite the growth in the availability of sequence data for honey bees, the phylogeny of the species remains a subject of controversy. Most notably, the geographic origin of honey bees is uncertain, as are the relationships among its constituent lineages and subspecies. We aim to infer the evolutionary and biogeographical history of the honey bee from mitochondrial genomes. Here we analyse the full mitochondrial genomes of 18 A. mellifera subspecies, belonging to all major lineages, using a range of gene sampling strategies and inference models to identify factors that may have contributed to the recovery of incongruent results in previous studies. Our analyses support a northern African or Middle Eastern origin of A. mellifera. We show that the previously suggested European and Afrotropical cradles of honey bees are the result of phylogenetic error. Monophyly of the M, C, and O lineages is strongly supported, but the A lineage appears paraphyletic. A. mellifera colonised Europe through at least two pathways, across the Strait of Gibraltar and via Asia Minor.

Osteostracans are the closest jawless relatives of jawed vertebrates, informing the gradual assembly of the vertebrate mineralised skeleton. Conflicting interpretations of their dermal skeletal histology arise from failure to account for topological variation, obscuring their significance in elucidating vertebrate skeletal evolution. To resolve this, we characterize the cranial and trunk dermal skeleton of a single individual of Tremataspis mammillata (Osteostraci, Thyestiida) at submicron resolution using synchrotron‐ and computed‐ tomography. Our results show that the architecture of the Tremataspis dermal skeleton is, for the most part, conserved over the skeleton and is broadly consistent with previous histological hypotheses based on 2D thin section study. We resolve debate over the homology of the basal layer, identifying it as osteogenic acellular isopedin rather than odontogenic elasmodine or metaplastic ossification of the stratum compactum of the dermis. We find topological variation between all dermal skeletal elements studied, and particularly between the cranial and postcranial dermal skeleton. This variation can be largely explained by reduction in differentiation due to geometric constraints imposed within smaller skeletal elements, such as scales. Our description of the dermal skeleton of Tremataspis mammillata provides a foundation for interpreting data from cursory topological samples of dermal skeletal diversity obtained in other osteostracans. This reveals general aspects of histological structure that must be ancestral for osteostracans and, likely, ancestral jawed vertebrates. Finally, we draw the distinction between hypotheses and descriptions in palaeohistology.

The distinctly non-random diversity of organis- mal form manifests itself in discrete clusters of taxa that share a common body plan. As a result, analyses of disparity require a scalable comparative framework. The difficulties of applying geometric morphometrics to disparity analyses of groups with vastly divergent body plans are overcome partly by the use of cladistic characters. Character-based disparity analyses have become increasingly popular, but it is not clear how they are affected by character coding strategies or revi- sions of primary homology statements. Indeed, whether cla- distic and morphometric data capture similar patterns of morphological variation remains a moot point. To address this issue, we employ both cladistic and geometric morpho- metric data in an exploratory study of disparity focussing on caecilian amphibians. Our results show no impact on relative intertaxon distances when different coding strategies for cla- distic characters were used or when revised concepts of homology were considered. In all instances, we found no sta- tistically significant difference between pairwise Euclidean and Procrustes distances, although the strength of the corre- lation among distance matrices varied. This suggests that cla- distic and geometric morphometric data appear to summarize morphological variation in comparable ways. Our results support the use of cladistic data for characteriz- ing organismal disparity.

Morphological variation (disparity) tends to be evaluated through two non-mutually exclusive approaches: (i) quantitatively, through geometric morphometrics, and (ii) in terms of discrete, ‘cladistic’, or categorical characters. Uncertainty over the comparability of these approaches diminishes the potential to obtain nomothetic insights into the evolution of morphological disparity, and the few benchmarking studies conducted so far show contrasting results. Here, we apply both approaches to characterising morphology in the stem-gnathostome vertebrate clade Osteostraci, in order to assess congruence between these alternative methods as well as to explore the evolutionary patterns of the group in terms of temporal disparity and the influence of phylogenetic relationships and habitat on morphospace occupation. Our results suggest that both approaches yield similar results in morphospace occupation and clustering, but also some differences indicating that these metrics may capture different aspects of morphology. Phylomorphospaces reveal important convergence towards a generalised ‘horseshoe’-shaped cranial morphology and two strong branching trends involving different major groups of osteostracans (benneviaspidids and thyestiids), which probably reflect adaptations to different lifestyles. Temporal patterns of disparity recorded by categorical and morphometric approaches differ considerably, capturing disparity maxima at very different times of the evolutionary history of the group. Disparity patterns recorded by the categorical approach parallel taxonomic diversity dynamics, likely reflecting a bias in facies representation rather than a real biological signal. This work provides evidence supporting that categorical and continuous data do not always capture morphological disparity in equivalent ways and that discrepancies reflect differences in the potential of each data type for characterizing more or less inclusive aspects of overall phenotype.

Stick and leaf insects (Phasmatodea) are a distinctive insect order whose members are characterized by mimicking various plant tissues such as twigs, foliage and bark. Unfortunately, the phylogenetic relationships among phasmatodean subfamilies and the timescale of their evolution remain uncertain. Recent molecular clock analyses have suggested a Cretaceous–Palaeogene origin of crown Phasmatodea and a subsequent Cenozoic radiation, contrasting with fossil evidence. Here, we analysed transcriptomic data from a broad diversity of phasmatodeans and, combined with the assembly of a new suite of fossil calibrations, we elucidate the evolutionary history of stick and leaf insects. Our results differ from recent studies in the position of the leaf insects (Phylliinae), which are recovered as sister to a clade comprising Clitumninae, Lancerocercata, Lonchodinae, Necrosciinae andXenophasmina. We recover a Permian to Triassic origin of crown Phasmatodea coinciding with the radiation of early insectivorous parareptiles, amphibians and synapsids. Aschiphasmatinae and Neophasmatodea diverged in the Jurassic–Early Cretaceous. A second spur in origination occurred in the Late Cretaceous, coinciding with the Cretaceous Terrestrial Revolution, and was probably driven by visual predators such as stem birds (Enantiornithes) and the radiation of angiosperms.