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The iconic marine raptorial predators Ichthyosauria and Eosauropterygia co-existed in the same ecosystems throughout most of the Mesozoic Era, facing similar evolutionary pressures and environmental perturbations. Both groups seemingly went through a massive macroevolutionary bottleneck across the Triassic-Jurassic (T/J) transition that greatly reduced their morphological diversity, leaving pelagic lineages as the only survivors. However, analyses of marine reptile disparity across the T/J transition have usually employed coarse morphological and temporal data. We comprehensively compare the evolution of ichthyosaurian and eosauropterygian morphology and body size across the Middle Triassic to Early Jurassic interval and find contrasting macroevolutionary patterns. The ecomorphospace of eosauropterygians predominantly reflects a strong phylogenetic signal, resulting in the clustering of three clades with clearly distinct craniodental phenotypes, suggesting "leaps" toward novel feeding ecologies. Ichthyosaurian diversification lacks a discernible evolutionary trend, as we find evidence for a wide overlap of craniodental morphologies between Triassic and Early Jurassic forms. The temporal evolution of ecomorphological disparity, fin shape and body size of eosauropterygians and ichthyosaurians during the Late Triassic does not support the hypothesis of an abrupt macroevolutionary bottleneck near the T/J transition. Rather, an important turnover event should be sought earlier, during times of rapid sea level falls., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE). All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Quantitative genetic theory on multivariate character evolution predicts that a population's response to directional selection is biased towards the major axis of the genetic covariance matrix G-the so-called genetic line of least resistance. Inferences on the genetic constraints in this sense have traditionally been made by measuring the angle of deviation of evolutionary trajectories from the major axis or, more recently, by calculating the amount of genetic variance-the Hansen-Houle evolvability-available along the trajectories. However, there have not been clear practical guidelines on how these quantities can be interpreted, especially in a high-dimensional space. This study summarizes pertinent distribution theories for relevant quantities, pointing out that they can be written as ratios of quadratic forms in evolutionary trajectory vectors by taking G as a parameter. For example, a beta distribution with appropriate parameters can be used as a null distribution for the squared cosine of the angle of deviation from a major axis or subspace. More general cases can be handled with the probability distribution of ratios of quadratic forms in normal variables. Apart from its use in hypothesis testing, this latter approach could potentially be used as a heuristic tool for looking into various selection scenarios, like directional and/or correlated selection, as parameterized with the mean and covariance of selection gradients., (© The Author(s) 2024. Published by Oxford University Press on behalf of the European Society of Evolutionary Biology.)

A dearth of Mesozoic-aged, three-dimensional fossils hinders understanding of the origin of the distinctive skull and brain of modern (crown) birds 1 . Here we report Navaornis hestiae gen. et sp. nov., an exquisitely preserved fossil species from the Late Cretaceous of Brazil. The skull of Navaornis is toothless and large-eyed, with a vaulted cranium closely resembling the condition in crown birds; however, phylogenetic analyses recover Navaornis in Enantiornithes, a highly diverse clade of Mesozoic stem birds. Despite an overall geometry quantitatively indistinguishable from crown birds, the skull of Navaornis retains numerous plesiomorphies including a maxilla-dominated rostrum, an akinetic palate, a diapsid temporal configuration, a small cerebellum and a weakly expanded telencephalon. These archaic neurocranial traits are combined with a crown bird-like degree of brain flexion and a bony labyrinth comparable in shape to those of many crown birds but substantially larger. Altogether, the emergent cranial geometry of Navaornis shows an unprecedented degree of similarity between crown birds and enantiornithines, groups last sharing a common ancestor more than 130 million years ago 2 . Navaornis provides long-sought insight into the detailed cranial and endocranial morphology of stem birds phylogenetically crownward of Archaeopteryx, clarifying the pattern and timing by which the distinctive neuroanatomy of living birds was assembled., Competing Interests: Competing interests The authors declare no competing interests., (© 2024. The Author(s).)

Birds' bills are their main tactile interface with the outside world. Tactile bill-tip organs associated with specialized foraging techniques are present in several bird groups, yet remain understudied in most clades. One example is Austrodyptornithes, the major seabird clade uniting Procellariiformes (albatrosses and petrels) and Sphenisciformes (penguins). Here, we describe the mechanoreceptor arrangement and neurovascular anatomy in the premaxillae of Austrodyptornithes. Using a wide phylogenetic sample of extant birds (361 species), we show that albatrosses and penguins exhibit complex tactile bill-tip anatomies, comparable to birds with known bill-tip organs, despite not being known to use tactile foraging. Petrels (Procellariidae, Hydrobatidae and Oceanitidae) lack these morphologies, indicating an evolutionary transition in bill-tip mechanosensitivity within Procellariiformes. The bill-tip organ in Austrodyptornithes may be functionally related to nocturnal foraging and prey detection under water, or courtship displays involving tactile stimulation of the bill. Alternatively, these organs may be vestigial as is likely the case in most palaeognaths (e.g. ostriches and emu). Ancestral state reconstructions fail to reject the hypothesis that the last common ancestor of Austrodyptornithes had a bill-tip organ; thus, tactile foraging may be ancestral for this major extant clade, perhaps retained from a deeper point in crown bird evolutionary history.

Lithornithidae, an assemblage of volant Palaeogene fossil birds, provide our clearest insights into the early evolutionary history of Palaeognathae, the clade that today includes the flightless ratites and volant tinamous. The neotype specimen of Lithornis vulturinus, from the early Eocene (approximately 53 million years ago) of Europe, includes a partial neurocranium that has never been thoroughly investigated. Here, we describe these cranial remains including the nearly complete digital endocasts of the brain and bony labyrinth. The telencephalon of Lithornis is expanded and its optic lobes are ventrally shifted, as is typical for crown birds. The foramen magnum is positioned caudally, rather than flexed ventrally as in some crown birds, with the optic lobes, cerebellum, and foramen magnum shifted further ventrally. The overall brain shape is similar to that of tinamous, the only extant clade of flying palaeognaths, suggesting that several aspects of tinamou neuroanatomy may have been evolutionarily conserved since at least the early Cenozoic. The estimated ratio of the optic lobe's surface area relative to the total brain suggests a diurnal ecology. Lithornis may provide the clearest insights to date into the neuroanatomy of the ancestral crown bird, combining an ancestrally unflexed brain with a caudally oriented connection with the spinal cord, a moderately enlarged telencephalon, and ventrally shifted, enlarged optic lobes., (© 2024 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

Estimates for animal antiquity exhibit a significant disconnect between those from molecular clocks, which indicate crown animals evolved ∼800 million years ago (Ma), and those from the fossil record, which extends only ∼574 Ma. Taphonomy is often held culpable: early animals were too small/soft/fragile to fossilise, or the circumstances that preserve them were uncommon in the early Neoproterozoic. We assess this idea by comparing Neoproterozoic fossilisation processes with those of the Cambrian and its abundant animal fossils. Cambrian Burgess Shale-type (BST) preservation captures animals in mudstones showing a narrow range of mineralogies; yet, fossiliferous Neoproterozoic mudstones rarely share the same mineralogy. Animal fossils are absent where BST preservation occurs in deposits ≥789 Ma, suggesting a soft maximum constraint on animal antiquity., Competing Interests: Declaration of interests The authors have no interests to declare., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Skeletal pneumaticity is a key feature of extant avian structure and biology, which first evolved among the non-flying archosaurian ancestors of birds. The widespread presence of air-filled bones across the postcranial skeleton is unique to birds among living vertebrates, but the true extent of skeletal pneumaticity has never been quantitatively investigated-hindering fundamental insights into the evolution of this key avian feature. Here, we use microCT scans of fresh, frozen birds to directly quantify the fraction of humerus volume occupied by air across a phylogenetically diverse taxon sample to test longstanding hypotheses regarding the evolution and function of avian skeletal pneumatization. Among other insights, we document weak positive allometry of internal air volume with humeral size among pneumatized humeri and provide strong support that humeral size, body mass, aquatic diving, and the presence or absence of pneumaticity all have independent effects on cortical bone thickness. Our quantitative evaluation of humeral pneumaticity across extant avian phylogeny sheds new light on the evolution and ontogenetic progression of an important aspect of avian skeletal architecture, and suggests that the last common ancestor of crown birds possessed a highly pneumatized humerus.

Characterizing how variation in the tempo and mode of evolution has structured the phenotypic diversity of extant species is a central goal of macroevolution 1-3 . However, studies are typically limited to a handful of traits 4-6 , providing incomplete information. We analyse morphological diversification in living birds, an ecologically diverse group 7 , documenting structural scales from 'pan-skeletal' proportions down to the localized three-dimensional shape changes of individual bones. We find substantial variation in evolutionary modes among avian subgroups and among skeletal parts, indicating widespread mosaicism and possible differences in the structure of the macroevolutionary landscape across Earth's main environments. Water-linked groups, especially Aequorlitornithes (waterbirds), have repeatedly explored a large portion of their total morphospace, emphasizing variation in body proportions and in the shape of bones close to the body core, which are functionally related to the mechanics of locomotion 8 . By contrast, landbirds (Inopinaves) evolved distinct, group-specific body forms early in the aftermath of the K-Pg and subsequently emphasized local shape variation, especially in the head and distal limb bones, which interact more directly with the environment. Passerines, which comprise more than half of all bird species, show a conservative evolutionary dynamic that resulted in low disparity across all skeletal parts. Evidence for early establishment of the morphospace of living birds is clear for some skeletal parts, including beaks and the combined skeletal morphology. However, we find little evidence for early partitioning of that morphospace, contrary to more specific predictions of 'niche-filling' models 1,9 . Nevertheless, early divergence among broad environmental types may have caused an early divergence of evolutionary modes, suggesting an important role for environmental divergence in structuring the radiation of crown-group birds., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

The origin of amniotes 320 million years ago signaled independence from water in vertebrates and was closely followed by divergences within the mammal and reptile stem lineages (Synapsida and Reptilia). Early members of both groups had highly similar morphologies, being superficially "lizard-like" forms with many plesiomorphies. However, the extent to which they might have exhibited divergent patterns of evolutionary change, with the potential to explain the large biological differences between their living members, is unresolved. We use a new, comprehensive phylogenetic dataset to quantify variation in rates and constraints of morphological evolution among Carboniferous-early Permian amniotes. We find evidence for an early burst of evolutionary rates, resulting in the early origins of morphologically distinctive subgroups that mostly persisted through the Cisuralian. Rates declined substantially through time, especially in reptiles. Early reptile evolution was also more constrained compared with early synapsids, exploring a more limited character state space. Postcranial innovation in particular was important in early synapsids, potentially related to their early origins of large body size. In contrast, early reptiles predominantly varied the temporal region, suggesting disparity in skull and jaw kinematics, and foreshadowing the variability of cranial biomechanics seen in reptiles today. Our results demonstrate that synapsids and reptiles underwent an early divergence of macroevolutionary patterns. This laid the foundation for subsequent evolutionary events and may be critical in understanding the substantial differences between mammals and reptiles today. Potential explanations include an early divergence of developmental processes or of ecological factors, warranting cross-disciplinary investigation. [Amniote; body size; constraint; phylogeny; rate.]., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society of Systematic Biologists.)

Parallelism between evolutionary trajectories in a trait space is often seen as evidence for repeatability of phenotypic evolution, and angles between trajectories play a pivotal role in the analysis of parallelism. However, properties of angles in multidimensional spaces have not been widely appreciated by biologists. To remedy this situation, this study provides a brief overview on geometric and statistical aspects of angles in multidimensional spaces. Under the null hypothesis that trajectory vectors have no preferred directions (i.e. uniform distribution on hypersphere), the angle between two independent vectors is concentrated around the right angle, with a more pronounced peak in a higher-dimensional space. This probability distribution is closely related to t - and beta distributions, which can be used for testing the null hypothesis concerning a pair of trajectories. A recently proposed method with eigenanalysis of a vector correlation matrix can be connected to the test of no correlation or concentration of multiple vectors, for which simple test procedures are available in the statistical literature. Concentration of vectors can also be examined by tools of directional statistics such as the Rayleigh test. These frameworks provide biologists with baselines to make statistically justified inferences for (non)parallel evolution.

Birds exhibit an enormous variety of beak shapes. Such remarkable variation, however, has distracted research from other important aspects of their skull evolution, the nature of which has been little explored. Key aspects of avian skull variation appear to be qualitatively similar to those of mammals, encompassing variation in the degree of cranial vaulting, cranial base flexure, and the proportions and orientations of the occipital and facial regions. The evolution of these traits has been studied intensively in mammals under the Spatial Packing Hypothesis (SPH), an architectural constraint so-called because the general anatomical organization and development of such skull parts makes them evolve predictably in response to changes in relative brain size. Such SPH predictions account for the different appearances of skull configurations across species, either in having longer or shorter faces, and caudally or ventrally oriented occiputs, respectively. This pattern has been morphometrically and experimentally proven in mammals but has not been examined in birds or other tetrapods, and so its generality remains unknown. We explored the SPH in an interspecific sample of birds using three-dimensional geometric morphometrics. Our results show that the dominant trend of evolutionary variation in the skull of crown-group birds can be predicted by the SPH, involving concomitant changes in the face, the cranial vault and the basicranium, and with striking similarities to craniofacial variation among mammals. Although craniofacial variation is significantly affected by allometry, these allometric effects are independent of the influence of the SPH on skull morphology, as are any effects of volumetric encephalization. Our results, therefore, validate the hypothesis that a general architectural constraint underlies skull homoplasy evolution of cranial morphology among avian clades, and possibly between birds and mammals, but they downplay encephalization and allometry as the only factors involved., (© 2021 Anatomical Society.)

Analysis of trait covariation plays a pivotal role in the study of phenotypic evolution. The magnitude of covariation is often quantified with statistics based on dispersion of eigenvalues of a covariance or correlation matrix-eigenvalue dispersion indices. This study clarifies the statistical justifications of these statistics and elaborates on their sampling properties. The relative eigenvalue variance of a covariance matrix is known in the statistical literature a test statistic for sphericity, and thus is an appropriate measure of eccentricity of variation. The same of a correlation matrix is equal to the average squared correlation, which has a straightforward interpretation as a measure of integration. Here, expressions for the mean and variance of these statistics are analytically derived under multivariate normality, clarifying the effects of sample size N, number of variables p, and parameters on sampling bias and error. Simulations confirm that approximations involved are reasonably accurate with a moderate sample size (N ≥ 16-64). Importantly, sampling properties of these indices are not adversely affected by a high p:N ratio, promising their utility in high-dimensional phenotypic analyses. They can furthermore be applied to shape variables and phylogenetically structured data with appropriate modifications., (© 2021 The Authors. Evolution © 2021 The Society for the Study of Evolution.)

Relative brain sizes in birds can rival those of primates, but large-scale patterns and drivers of avian brain evolution remain elusive. Here, we explore the evolution of the fundamental brain-body scaling relationship across the origin and evolution of birds. Using a comprehensive dataset sampling> 2,000 modern birds, fossil birds, and theropod dinosaurs, we infer patterns of brain-body co-variation in deep time. Our study confirms that no significant increase in relative brain size accompanied the trend toward miniaturization or evolution of flight during the theropod-bird transition. Critically, however, theropods and basal birds show weaker integration between brain size and body size, allowing for rapid changes in the brain-body relationship that set the stage for dramatic shifts in early crown birds. We infer that major shifts occurred rapidly in the aftermath of the Cretaceous-Paleogene mass extinction within Neoaves, in which multiple clades achieved higher relative brain sizes because of a reduction in body size. Parrots and corvids achieved the largest brains observed in birds via markedly different patterns. Parrots primarily reduced their body size, whereas corvids increased body and brain size simultaneously (with rates of brain size evolution outpacing rates of body size evolution). Collectively, these patterns suggest that an early adaptive radiation in brain size laid the foundation for subsequent selection and stabilization., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Fossils of the Ediacaran macrobiota (∼571-539 mya) record phylogenetically diverse marine palaeocommunities, including early animals, which pre-date the "Cambrian Explosion" [1-4]. Benthic forms with a frondose gross morphology, assigned to the morphogroups Rangeomorpha [5] and Frondomorpha (see also Arboreomorpha) [6-8], are among the most temporally wide-ranging and environmentally tolerant members of the Ediacaran macrobiota [6] and dominated deep-marine ecosystems ∼571-560 mya [9-11]. Investigations into the morphology [12-14], palaeoecology [10, 15, 16], reproductive strategies [17, 18], feeding methods [9, 19], and morphogenesis of frondose taxa together constrain their phylogenetic position to the metazoan (for Rangeomorpha) or eumetazoan (e.g., Arborea) total groups [14, 20], but tighter constraint is currently lacking. Here, we describe fossils of abundant filamentous organic structures preserved among frond-dominated fossil assemblages in Newfoundland (Canada). The filaments constitute a prominent component of the ecosystems, and exhibit clear physical associations with at least seven frondose taxa. Individual specimens of one uniterminal rangeomorph taxon appear to be directly connected by filaments across distances of centimeters to meters. Such physical linkages are interpreted to reflect evidence for stolonic connections: a conclusion with potential implications for the phylogenetic placement and palaeoecology of frondose organisms. Consideration of extant stoloniferous organisms suggests that Ediacaran frondose taxa were likely clonal and resurrects the possibility that they may have been colonial (e.g., [21, 22]). VIDEO ABSTRACT., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The 'Cambrian Explosion' describes the rapid increase in animal diversity and abundance, as manifest in the fossil record, between ~540 and 520 million years ago (Ma). This event, however, is nested within a far more ancient record of macrofossils extending at least into the late Ediacaran at ~571 Ma. The evolutionary events documented during the Ediacaran-Cambrian interval coincide with geochemical evidence for the modernisation of Earth's biogeochemical cycles. Holistic integration of fossil and geochemical records leads us to challenge the notion that the Ediacaran and Cambrian worlds were markedly distinct, and places biotic and environmental change within a longer-term narrative. We propose that the evolution of metazoans may have been facilitated by a series of dynamic and global changes in redox conditions and nutrient supply, which, potentially together with biotic feedbacks, enabled turnover events that sustained multiple phases of radiation. We argue that early metazoan diversification should be recast as a series of successive, transitional radiations that extended from the late Ediacaran and continued through the early Palaeozoic. We conclude that while the Cambrian Explosion represents a radiation of crown-group bilaterians, it was simply one phase amongst several metazoan radiations, some older and some younger.

Cambrian annelids are strikingly diverse and reveal important details of annelid character acquisition. Their contribution, however, to a wider understanding of the evolution of the trochozoans (encompassing the annelids as well as such groups as the brachiopods and molluscs) remains limited. Thus the early annelids had been linked to a variety of cataphract Cambrian metazoans, notably Wiwaxia and the halkieriids, but recent work assigns such fossils to stem-group molluscs. Here we report two new annelids from the Lower Cambrian Chengjiang Lagerstätte, South China. Ipoliknus avitus n. gen., n. sp. is biramous with neurochaetae and notochaetae, but significantly also bears dorsal spinose sclerites and dorso-lateral dentate sclerites. Adelochaeta sinensis n. gen., n. sp. is unique amongst Cambrian polychaetes in possessing the rod-like supports of the parapodia known as aciculae. This supports phylogenetic placement of Adelochaeta as sister to some more derived aciculate Palaeozoic taxa, but in contrast Ipoliknus is recovered as the most basal of the stem-group annelids. Sclerites and chaetae of I. avitus are interpreted respectively as the remnants and derivatives of a once more extensive cataphract covering that was a characteristic of more primitive trochozoans. The two sets of chaetae (noto- and neurochaetae) and two sets of sclerites (spinose and dentate) suggest that in a pre-annelid an earlier and more complete scleritome may have consisted of four zones of sclerites. Other cataphract taxa from the Lower Palaeozoic show a variety of scleritome configurations but establishing direct links with such basal annelids as Ipoliknus at present must remain conjectural.

Through elasmobranch (sharks and rays) evolutionary history, gigantism evolved multiple times in phylogenetically distant species, some of which are now extinct. Interestingly, the world's largest elasmobranchs display two specializations found never to overlap: filter feeding and mesothermy. The contrasting lifestyles of elasmobranch giants provide an ideal case study to elucidate the evolutionary pathways leading to gigantism in the oceans. Here, we applied a phylogenetic approach to a global dataset of 459 taxa to study the evolution of elasmobranch gigantism. We found that filter feeders and mesotherms deviate from general relationships between trophic level and body size, and exhibit significantly larger sizes than ectothermic-macropredators. We confirm that filter feeding arose multiple times during the Paleogene, and suggest the possibility of a single origin of mesothermy in the Cretaceous. Together, our results elucidate two main evolutionary pathways that enable gigantism: mesothermic and filter feeding. These pathways were followed by ancestrally large clades and facilitated extreme sizes through specializations for enhancing prey intake. Although a negligible percentage of ectothermic-macropredators reach gigantic sizes, these species lack such specializations and are correspondingly constrained to the lower limits of gigantism. Importantly, the very adaptive strategies that enabled the evolution of the largest sharks can also confer high extinction susceptibility., (© 2019 The Author(s). Evolution © 2019 The Society for the Study of Evolution.)

New fossils help pinpoint when some birds started relying on a seed-based diet and reveal that disparate bill shapes evolved repeatedly throughout bird evolutionary history., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The first known diverse, complex, macroscopic benthic marine ecosystems (late Ediacaran, ca. 571-541 Ma) were dominated by the Rangeomorpha, an enigmatic group of extinct frondose eukaryotes that are candidate early metazoans [1, 2]. The group is characterized by a self-similar branching architecture that was most likely optimized for exchange, but nearly every other aspect of their biology is contentious [2-4]. We report locally enhanced, aberrant growth ("eccentric branching") in a stalked, multifoliate rangeomorph-Hylaecullulus fordi n. gen., n. sp.-from Charnwood Forest (UK), confirming the presence of true biological modularity within the group. Random branches achieve unusually large proportions and mimic the architecture of their parent branch, rather than that of their neighbors (the norm). Their locations indicate exceptional growth at existing loci, rather than insertion at new sites. Analogous overcompensatory branching in extant modular organisms requires the capacity to orchestrate growth at specific sites and occurs most frequently in response to damage or environmental stress, allowing regeneration toward optimum morphology (e.g., [5-7]). Its presence in rangeomorphs indicates a hitherto unappreciated level of control to their growth plan, a previously unrecognized form of morphological plasticity within the group, and an ability to actively respond to external physical stimuli. The trait would have afforded rangeomorphs resilience to fouling and abrasion, partially accounting for their wide environmental tolerance, and may have pre-adapted them to withstand predation, weakening this argument for their extinction. Our findings highlight that multiple, phylogenetically disparate clades first achieved large size through modularity., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The evolutionary events during the Ediacaran-Cambrian transition (~541 Myr ago) are unparalleled in Earth history. The fossil record suggests that most extant animal phyla appeared in a geologically brief interval, with the oldest unequivocal bilaterian body fossils found in the Early Cambrian. Molecular clocks and biomarkers provide independent estimates for the timing of animal origins, and both suggest a cryptic Neoproterozoic history for Metazoa that extends considerably beyond the Cambrian fossil record. We report an assemblage of ichnofossils from Ediacaran-Cambrian siltstones in Brazil, alongside U-Pb radioisotopic dates that constrain the age of the oldest specimens to 555-542 Myr. X-ray microtomography reveals three-dimensionally preserved traces ranging from 50 to 600 μm in diameter, indicative of small-bodied, meiofaunal tracemakers. Burrow morphologies suggest they were created by a nematoid-like organism that used undulating locomotion to move through the sediment. This assemblage demonstrates animal-sediment interactions in the latest Ediacaran period, and provides the oldest known fossil evidence for meiofaunal bilaterians. Our discovery highlights meiofaunal ichnofossils as a hitherto unexplored window for tracking animal evolution in deep time, and reveals that both meiofaunal and macrofaunal bilaterians began to explore infaunal niches during the late Ediacaran.

The late Ediacaran soft-bodied macroorganism Dickinsonia (age range approx. 560-550 Ma) has often been interpreted as an early animal, and is increasingly invoked in debate on the evolutionary assembly of eumetazoan body plans. However, conclusive positive evidence in support of such a phylogenetic affinity has not been forthcoming. Here we subject a collection of Dickinsonia specimens interpreted to represent multiple ontogenetic stages to a novel, quantitative method for studying growth and development in organisms with an iterative body plan. Our study demonstrates that Dickinsonia grew via pre-terminal 'deltoidal' insertion and inflation of constructional units, followed by a later inflation-dominated phase of growth. This growth model is contrary to the widely held assumption that Dickinsonia grew via terminal addition of units at the end of the organism bearing the smallest units. When considered alongside morphological and behavioural attributes, our developmental data phylogenetically constrain Dickinsonia to the Metazoa, specifically the Eumetazoa plus Placozoa total group. Our findings have implications for the use of Dickinsonia in developmental debates surrounding the metazoan acquisition of axis specification and metamerism., (© 2017 The Author(s).)

Macroscale rangeomorph fossils, with characteristic branching fronds, appear (571 Myr ago) after the Gaskiers glaciation (580 Myr ago). However, biological mechanisms of size growth and potential connections to ocean geochemistry were untested. Using micro-computerized tomography and photographic measurements, alongside mathematical and computer models, we demonstrate that growth of rangeomorph branch internodes declined as their relative surface area decreased. This suggests that frond size and shape were directly responsive to nutrient uptake.

Morphospaces are spatial depictions of morphological variation among biological forms that have become an integral part of the analytical toolkit of evolutionary biologists and palaeobiologists. Nevertheless, the term morphospace brings together a great variety of spaces with different geometries. In particular, many morphospaces lack the metric properties underlying the notions of distance and direction, which are, however, central to the analysis of morphological differences and evolutionary transitions. The problem is illustrated here with the iconic morphospace of coiled shells implemented by Raup 50 years ago. The model, which allows the description of shell coiling geometry of various invertebrate taxa, is a seminal reference in theoretical morphology and morphospace theory, but also a morphometric framework frequently used in empirical studies, particularly of ammonoids. Because of the definition of its underlying parameters, Raup's morphospace does not possess a Euclidean structure and a meaningful interpretation of the spread and spacing of taxa within it is not guaranteed. Focusing on the region of the morphospace occupied by most ammonoids, I detail a landmark-based morphospace circumventing this problem and built from the same input measurements required for the calculation of Raup's parameters. From simulations and a reanalysis of Palaeozoic ammonoid shell disparity, the properties of these morphospaces are compared and their algebraic and geometric relationships highlighted. While Raup's model remains a valuable tool for describing ammonoid shells and relating their shapes to the coiling process, it is demonstrated that quantitative analyses of morphological patterns should be carried out within the landmark-based framework. Beyond this specific case, the increasing use and diversity of morphospaces in evolutionary morphology call for caution when interpreting patterns and comparing results drawn from different types of morphospaces., (© 2016 Cambridge Philosophical Society.)

Deuterostomes include the group we belong to (vertebrates) as well as an array of disparate forms that include echinoderms, hemichordates and more problematic groups such as vetulicolians and vetulocystids. The Cambrian fossil record is well-populated with representative examples, but possible intermediates are controversial and the nature of the original deuterostome remains idealized. Here we report millimetric fossils, Saccorhytus coronarius nov. gen., nov. sp., from an Orsten-like Lagerstätte from the earliest Cambrian period of South China, which stratigraphically are amongst the earliest of deuterostomes. The bag-like body bears a prominent mouth and associated folds, and behind them up to four conical openings on either side of the body as well as possible sensory structures. An anus may have been absent, and correspondingly the lateral openings probably served to expel water and waste material. This new form has similarities to both the vetulicolians and vetulocystids and collectively these findings suggest that a key step in deuterostome evolution was the development of lateral openings that subsequently were co-opted as pharyngeal gills. Depending on its exact phylogenetic position, the meiofaunal habit of Saccorhytus may help to explain the major gap between divergence times seen in the fossil record and estimates based on molecular clocks.

The evolution of vertebral fusion is a poorly understood phenomenon that results in the loss of mobility between sequential vertebrae. Non-pathological fusion of the anterior cervical vertebrae has evolved independently in numerous extant and extinct mammals and reptiles, suggesting that the formation of a 'syncervical' is an adaptation that arose to confer biomechanical advantage(s) in these lineages. We review syncervical anatomy and evolution in a broad phylogenetic context for the first time and provide a comprehensive summary of proposed adaptive hypotheses. The syncervical generally consists of two vertebrae (e.g. hornbills, porcupines, dolphins) but can include fusion of seven cervical vertebrae in some cetaceans. Based on the ecologies of taxa with this trait, cervical fusion most often occurs in fossorial and pelagic taxa. In fossorial taxa, the syncervical likely increases the out-lever force during head-lift digging. In cetaceans and ricochetal rodents, the syncervical may stabilize the head and neck during locomotion, although considerable variation exists in its composition without apparent variability in locomotion. Alternatively, the highly reduced cervical vertebral centra may require fusion to prevent mechanical failure of the vertebrae. In birds, the syncervical of hornbills may have evolved in response to their unique casque-butting behaviour, or due to increased head mass. The general correlation between ecological traits and the presence of a syncervical in extant taxa allows more accurate interpretation of extinct animals that also exhibit this unique trait. For example, syncervicals evolved independently in several groups of marine reptiles and may have functioned to stabilize the head at the craniocervical joint during pelagic locomotion, as in cetaceans. Overall, the origin and function of fused cervical vertebrae is poorly understood, emphasizing the need for future comparative biomechanical studies interpreted in an evolutionary context., (© 2016 Cambridge Philosophical Society.)

Australian spiny mountain crayfish (Euastacus, Parastacidae) and their ecotosymbiotic temnocephalan flatworms (Temnocephalida, Platyhelminthes) may have co-occurred and interacted through deep time, during a period of major environmental change. Therefore, reconstructing the history of their association is of evolutionary, ecological, and conservation significance. Here, time-calibrated Bayesian phylogenies of Euastacus species and their temnocephalans (Temnohaswellia and Temnosewellia) indicate near-synchronous diversifications from the Cretaceous. Statistically significant cophylogeny correlations between associated clades suggest linked evolutionary histories. However, there is a stronger signal of codivergence and greater host specificity in Temnosewellia, which co-occurs with Euastacus across its range. Phylogeography and analyses of evolutionary distinctiveness (ED) suggest that regional differences in the impact of climate warming and drying had major effects both on crayfish and associated temnocephalans. In particular, Euastacus and Temnosewellia show strong latitudinal gradients in ED and, conversely, in geographical range size, with the most distinctive, northern lineages facing the greatest risk of extinction. Therefore, environmental change has, in some cases, strengthened ecological and evolutionary associations, leaving host-specific temnocephalans vulnerable to coextinction with endangered hosts. Consequently, the extinction of all Euastacus species currently endangered (75%) predicts coextinction of approximately 60% of the studied temnocephalans, with greatest loss of the most evolutionarily distinctive lineages., (© 2016 The Author(s).)

The ever-increasing number of studies that address the origin and evolution of Euarthropoda - whose extant representatives include chelicerates, myriapods, crustaceans and hexapods - are gradually reaching a consensus with regard to the overall phylogenetic relationships of some of the earliest representatives of this phylum. The stem-lineage of Euarthropoda includes numerous forms that reflect the major morphological transition from a lobopodian-type to a completely arthrodized body organization. Several methods of classification that aim to reflect such a complex evolutionary history have been proposed as a consequence of this taxonomic diversity. Unfortunately, this has also led to a saturation of nomenclatural schemes, often in conflict with each other, some of which are incompatible with cladistic-based methodologies. Here, I review the convoluted terminology associated with the classification of stem-group Euarthropoda, and propose a synapomorphy-based distinction that allows 'lower stem-Euarthropoda' (e.g. lobopodians, radiodontans) to be separated from 'upper stem-Euarthropoda' (e.g. fuxianhuiids, Cambrian bivalved forms) in terms of the structural organization of the head region and other aspects of overall body architecture. The step-wise acquisition of morphological features associated with the origins of the crown-group indicate that the node defining upper stem-Euarthropoda is phylogenetically stable, and supported by numerous synapomorphic characters; these include the presence of a deutocerebral first appendage pair, multisegmented head region with one or more pairs of post-ocular differentiated limbs, complete body arthrodization, posterior-facing mouth associated with the hypostome/labrum complex, and post-oral biramous arthropodized appendages. The name 'Deuteropoda' nov. is proposed for the scion (monophyletic group including the crown-group and an extension of the stem-group) that comprises upper stem-Euarthropoda and Euarthropoda. A brief account of common terminological inaccuracies in recent palaeontological studies evinces the utility of Deuteropoda nov. as a reference point for discussing aspects of early euarthropod phylogeny., (© 2014 Cambridge Philosophical Society.)

Examples of long-term coevolution are rare among free-living organisms. Müllerian mimicry in Heliconius butterflies had been suggested as a key example of coevolution by early genetic studies. However, research over the last two decades has been dominated by the idea that the best-studied comimics, H. erato and H. melpomene, did not coevolve at all. Recently sequenced genes associated with wing color pattern phenotype offer a new opportunity to resolve this controversy. Here, we test the hypothesis of coevolution between H. erato and H. melpomene using Bayesian multilocus analysis of five color pattern genes and five neutral genetic markers. We first explore the extent of phylogenetic agreement versus conflict between the different genes. Coevolution is then tested against three aspects of the mimicry diversifications: phylogenetic branching patterns, divergence times, and, for the first time, phylogeographic histories. We show that all three lines of evidence are compatible with strict coevolution of the diverse mimicry wing patterns, contrary to some recent suggestions. Instead, these findings tally with a coevolutionary diversification driven primarily by the ecological force of Müllerian mimicry., (© 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.)

The Neoproterozoic era was arguably the most revolutionary in Earth history. Extending from 1000 to 541 million years ago, it stands at the intersection of the two great tracts of evolutionary time: on the one side, some three billion years of pervasively microbial 'Precambrian' life, and on the other the modern 'Phanerozoic' biosphere with its extraordinary diversity of large multicellular organisms. The disturbance doesn't stop here, however: over this same stretch of time the planet itself was in the throes of change. Tectonically, it saw major super-continental reconfigurations, climatically its deepest ever glacial freeze, and geochemically some of the most anomalous perturbations on record. What lies behind this dramatic convergence of biological and geological phenomena, and how exactly did it give rise to the curiously complex world that we now inhabit?, (Copyright © 2015 Elsevier Ltd. All rights reserved.)

We describe Collinsium ciliosum from the early Cambrian Xiaoshiba Lagerstätte in South China, an armored lobopodian with a remarkable degree of limb differentiation including a pair of antenna-like appendages, six pairs of elongate setiferous limbs for suspension feeding, and nine pairs of clawed annulated legs with an anchoring function. Collinsium belongs to a highly derived clade of lobopodians within stem group Onychophora, distinguished by a substantial dorsal armature of supernumerary and biomineralized spines (Family Luolishaniidae). As demonstrated here, luolishaniids display the highest degree of limb specialization among Paleozoic lobopodians, constitute more than one-third of the overall morphological disparity of stem group Onychophora, and are substantially more disparate than crown group representatives. Despite having higher disparity and appendage complexity than other lobopodians and extant velvet worms, the specialized mode of life embodied by luolishaniids became extinct during the Early Paleozoic. Collinsium and other superarmored lobopodians exploited a unique paleoecological niche during the Cambrian explosion.

The anterior cervical vertebrae form the skeletal connection between the cranial and postcranial skeletons in higher tetrapods. As a result, the morphology of the atlas-axis complex is likely to be shaped by selection pressures acting on either the head or neck. The neoceratopsian (Reptilia:Dinosauria) syncervical represents one of the most highly modified atlas-axis regions in vertebrates, being formed by the complete coalescence of the three most anterior cervical vertebrae. In ceratopsids, the syncervical has been hypothesized to be an adaptation to support a massive skull, or to act as a buttress during intraspecific head-to-head combat. Here, we test these functional/adaptive hypotheses within a phylogenetic framework and critically examine the previously proposed methods for quantifying relative head size in the fossil record for the first time. Results indicate that neither the evolution of cranial weaponry nor large head size correlates with the origin of cervical fusion in ceratopsians, and we, therefore, reject both adaptive hypotheses for the origin of the syncervical. Anterior cervical fusion has evolved independently in a number of amniote clades, and further research on extant groups with this peculiar anatomy is needed to understand the evolutionary basis for cervical fusion in Neoceratopsia., (© 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.)

Morphospaces are quantitative representations of phenotype space that are widely used in studies of morphological evolution. Do current conceptualizations of morphospaces, however, appropriately reflect the evolutionary dynamics of organisms depicted in these spaces? Most empirical morphospace studies implicitly consider variability of biological forms as isotropic, but such a view appears inadequate when the properties of development mediating phenotypic changes are considered. Here, a trilobite case study is used to visualize the constraints imposed by development on the accessibility structure of morphospace. Variability in the resultant morphospace is strongly anisotropic and reveals discordances between the apparent range of possible phenotypes and their actual accessibility. Homoplasy, directionality, and asymmetry of evolutionary transitions appear as natural consequences of anisotropic variability and point out the limitation of morphological distance for evolutionary inference. Measures of distance in morphospace should be used with considerable caution and must be complemented with developmentally meaningful measures of evolutionary accessibility., (© 2014 Wiley Periodicals, Inc.)

Unlabelled: •, Premise of the Study: Globally, the origins of xeromorphic traits in modern angiosperm lineages are obscure but are thought to be linked to the early Neogene onset of seasonally arid climates. Stomatal encryption is a xeromorphic trait that is prominent in Banksia, an archetypal genus centered in one of the world's most diverse ecosystems, the ancient infertile landscape of Mediterranean-climate southwestern Australia.•, Methods: We describe Banksia paleocrypta, a sclerophyllous species with encrypted stomata from silcretes of the Walebing and Kojonup regions of southwestern Australia dated as Late Eocene.•, Key Results: Banksia paleocrypta shows evidence of foliar xeromorphy ∼20 Ma before the widely accepted timing for the onset of aridity in Australia. Species of Banksia subgenus Banksia with very similar leaves are extant in southwestern Australia. The conditions required for silcrete formation infer fluctuating water tables and climatic seasonality in southwestern Australia in the Eocene, and seasonality is supported by the paucity of angiosperm closed-forest elements among the fossil taxa preserved with B. paleocrypta. However, climates in the region during the Eocene are unlikely to have experienced seasons as hot and dry as present-day summers.•, Conclusions: The presence of B. paleocrypta within the center of diversity of subgenus Banksia in edaphically ancient southwestern Australia is consistent with the continuous presence of this lineage in the region for ≥40 Ma, a testament to the success of increasingly xeromorphic traits in Banksia over an interval in which numerous other lineages became extinct., (© 2014 Botanical Society of America, Inc.)

Trilobites are typified by the behavioural and morphological ability to enrol their bodies, most probably as a defence mechanism against adverse environmental conditions or predators. Although most trilobites could enrol at least partially, there is uncertainty about whether olenellids-among the most phylogenetically and stratigraphically basal representatives-could perform this behaviour because of their poorly caudalized trunk and scarcity of coaptative devices. Here, we report complete-but not encapsulating-enrolment for the olenellid genus Mummaspis from the early Cambrian Mural Formation in Alberta, the earliest direct evidence of this strategy in the fossil record of polymerid trilobites. Complete enrolment in olenellids was achieved through a combination of ancestral morphological features, and thus provides new information on the character polarity associated with this key trilobite adaptation.

The Middle Cambrian Pikaia gracilens (Walcott) has an iconic position as a Cambrian chordate, but until now no detailed description has been available. Here on the basis of the 114 available specimens we review its anatomy, confirm its place in the chordates and explore with varying degrees of confidence its relationships to both extant and extinct chordates and other deuterostomes. The body of Pikaia is fusiform, laterally compressed and possesses about 100 myomeres. The head is small, bilobed and bears two narrow tentacles. There is no evidence for eyes. Apart from a thin dorsal fin (without finrays) and a series of at least nine bilaterally arranged appendages with possible pharyngeal pores at the anterior end, there are no other external features. In addition to the musculature the internal anatomy includes an alimentary canal, the anterior of which forms a prominent lenticular unit that is almost invariably preserved in positive relief. The cavity is interpreted as pharyngeal, implying that the mouth itself was almost terminal. The posterior extension of the gut is unclear although the anus appears to have been terminal. The most prominent internal structure is a reflectively preserved unit, possibly hollow, termed here the dorsal organ. Although formerly interpreted as a notochord its position and size make this less likely. Its original function remains uncertain, but it could have formed a storage organ. Ventral to the dorsal organ a narrower strand of tissue is interpreted as representing the nerve chord and notochord. In addition to these structures, there is also evidence for a vascular system, including a ventral blood vessel. The position of Pikaia in the chordates is largely based on the presence of sigmoidal myomeres, and the more tentative identification of a notochord. In many other respects, Pikaia differs from the expected nature of primitive chordates, especially as revealed in amphioxus and the Cambrian record (including Cathaymyrus, Haikouichthys, Metaspriggina, Myllokunmingia, and Zhongxiniscus). Whilst the possibility that Pikaia is simply convergent on the chordates cannot be dismissed, we prefer to build a scenario that regards Pikaia as the most stem-ward of the chordates with links to the phylogenetically controversial yunnanozoans. This hypothesis has implications for the evolution of the myomeres, notochord and gills. Finally, the wealth of material of Pikaia indicates that, although by definition there must be some sort of taphonomic imprint, the consistency of preservational details allows a reliable reconstruction of the anatomy and does not significantly erode phylogenetically relevant characters., (© 2012 The Authors. Biological Reviews © 2012 Cambridge Philosophical Society.)

Statistical models are helping palaeontologists to elucidate the history of biodiversity. Sampling standardization has been extensively applied to remedy the effects of uneven sampling in large datasets of fossil invertebrates. However, many vertebrate datasets are smaller, and the issue of uneven sampling has commonly been ignored, or approached using pairwise comparisons with a numerical proxy for sampling effort. Although most authors find a strong correlation between palaeodiversity and sampling proxies, weak correlation is recorded in some datasets. This has led several authors to conclude that uneven sampling does not influence our view of vertebrate macroevolution. We demonstrate that multi-variate regression models incorporating a model of underlying biological diversification, as well as a sampling proxy, fit observed sauropodomorph dinosaur palaeodiversity best. This bivariate model is a better fit than separate univariate models, and illustrates that observed palaeodiversity is a composite pattern, representing a biological signal overprinted by variation in sampling effort. Multi-variate models and other approaches that consider sampling as an essential component of palaeodiversity are central to gaining a more complete understanding of deep time vertebrate diversification.

Pneumatic (air-filled) postcranial bones are unique to birds among extant tetrapods. Unambiguous skeletal correlates of postcranial pneumaticity first appeared in the Late Triassic (approximately 210 million years ago), when they evolved independently in several groups of bird-line archosaurs (ornithodirans). These include the theropod dinosaurs (of which birds are extant representatives), the pterosaurs, and sauropodomorph dinosaurs. Postulated functions of skeletal pneumatisation include weight reduction in large-bodied or flying taxa, and density reduction resulting in energetic savings during foraging and locomotion. However, the influence of these hypotheses on the early evolution of pneumaticity has not been studied in detail previously. We review recent work on the significance of pneumaticity for understanding the biology of extinct ornithodirans, and present detailed new data on the proportion of the skeleton that was pneumatised in 131 non-avian theropods and Archaeopteryx. This includes all taxa known from significant postcranial remains. Pneumaticity of the cervical and anterior dorsal vertebrae occurred early in theropod evolution. This 'common pattern' was conserved on the line leading to birds, and is likely present in Archaeopteryx. Increases in skeletal pneumaticity occurred independently in as many as 12 lineages, highlighting a remarkably high number of parallel acquisitions of a bird-like feature among non-avian theropods. Using a quantitative comparative framework, we show that evolutionary increases in skeletal pneumaticity are significantly concentrated in lineages with large body size, suggesting that mass reduction in response to gravitational constraints at large body sizes influenced the early evolution of pneumaticity. However, the body size threshold for extensive pneumatisation is lower in theropod lineages more closely related to birds (maniraptorans). Thus, relaxation of the relationship between body size and pneumatisation preceded the origin of birds and cannot be explained as an adaptation for flight. We hypothesise that skeletal density modulation in small, non-volant, maniraptorans resulted in energetic savings as part of a multi-system response to increased metabolic demands. Acquisition of extensive postcranial pneumaticity in small-bodied maniraptorans may indicate avian-like high-performance endothermy., (© 2011 The Authors. Biological Reviews © 2011 Cambridge Philosophical Society.)

Animals do not just occupy the modern biosphere, they permeate its structure and define how it works. Their unique combination of organ-grade multicellularity, motility and heterotrophic habit makes them powerful geobiological agents, imposing myriad feedbacks on nutrient cycling, productivity and environment. Most significantly, animals have 'engineered' the biosphere over evolutionary time, forcing the diversification of, for example, phytoplankton, land plants, trophic structure, large body size, bioturbation, biomineralization and indeed the evolutionary process itself. This review surveys how animals contribute to the modern world and provides a basis for reconstructing ancient ecosystems. Earlier, less animal-influenced biospheres worked quite differently from the one currently occupied, with the Ediacaran-Cambrian radiation of organ-grade animals marking a fundamental shift in macroecological and macroevolutionary expression., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

The evolution of mammalian brain size is directly linked with the evolution of the brain's unique structure and performance. Both maternal life history investment traits and basal metabolic rate (BMR) correlate with relative brain size, but current hypotheses regarding the details of these relationships are based largely on placental mammals. Using encephalization quotients, partial correlation analyses, and bivariate regressions relating brain size to maternal investment times and BMR, we provide a direct quantitative comparison of brain size evolution in marsupials and placentals, whose reproduction and metabolism differ extensively. Our results show that the misconception that marsupials are systematically smaller-brained than placentals is driven by the inclusion of one large-brained placental clade, Primates. Marsupial and placental brain size partial correlations differ in that marsupials lack a partial correlation of BMR with brain size. This contradicts hypotheses stating that the maintenance of relatively larger brains requires higher BMRs. We suggest that a positive BMR-brain size correlation is a placental trait related to the intimate physiological contact between mother and offspring during gestation. Marsupials instead achieve brain sizes comparable to placentals through extended lactation. Comparison with avian brain evolution suggests that placental brain size should be constrained due to placentals' relative precociality, as has been hypothesized for precocial bird hatchlings. We propose that placentals circumvent this constraint because of their focus on gestation, as opposed to the marsupial emphasis on lactation. Marsupials represent a less constrained condition, demonstrating that hypotheses regarding placental brain size evolution cannot be generalized to all mammals.

Evolutionary biology rejoices in the diversity of life, but this comes at a cost: other than working in the common framework of neo-Darwinian evolution, specialists in, for example, diatoms and mammals have little to say to each other. Accordingly, their research tends to track the particularities and peculiarities of a given group and seldom enquires whether there are any wider or deeper sets of explanations. Here, I present evidence in support of the heterodox idea that evolution might look to a general theory that does more than serve as a tautology ('evolution explains evolution'). Specifically, I argue that far from its myriad of products being fortuitous and accidental, evolution is remarkably predictable. Thus, I urge a move away from the continuing obsession with Darwinian mechanisms, which are entirely uncontroversial. Rather, I emphasize why we should seek explanations for ubiquitous evolutionary convergence, as well as the emergence of complex integrated systems. At present, evolutionary theory seems to be akin to nineteenth-century physics, blissfully unaware of the imminent arrival of quantum mechanics and general relativity. Physics had its Newton, biology its Darwin: evolutionary biology now awaits its Einstein.

The very success of the Darwinian explanation, in not only demonstrating evolution from multiple lines of evidence but also in providing some plausible explanations, paradoxically seems to have served to have stifled explorations into other areas of investigation. The fact of evolution is now almost universally yoked to the assumption that its outcomes are random, trends are little more than drunkard's walks, and most evolutionary products are masterpieces of improvisation and far from perfect. But is this correct? Let us consider some alternatives. Is there evidence that evolution could in anyway be predictable? Can we identify alternative forms of biological organizations and if so how viable are they? Why are some molecules so extraordinarily versatile, while others can be spoken of as "molecules of choice"? How fortuitous are the major transitions in the history of life? What implications might this have for the Tree of Life? To what extent is evolutionary diversification constrained or facilitated by prior states? Are evolutionary outcomes merely sufficient or alternatively are they highly efficient, even superb? Here I argue that in sharp contradistinction to an orthodox Darwinian view, not only is evolution much more predictable than generally assumed but also investigation of its organizational substrates, including those of sensory systems, which indicates that it is possible to identify a predictability to the process and outcomes of evolution. If correct, the implications may be of some significance, not least in separating the unexceptional Darwinian mechanisms from underlying organizational principles, which may indicate evolutionary inevitabilities.

Halkieriids and wiwaxiids are cosmopolitan sclerite-bearing metazoans from the Lower and Middle Cambrian. Although they have similar scleritomes, their phylogenetic position is contested. A new scleritomous fossil from the Burgess Shale has the prominent anterior shell of the halkieriids but also bears wiwaxiid-like sclerites. This new fossil defines the monophyletic halwaxiids and indicates that they have a key place in early lophotrochozoan history.