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Observations of maxillary (upper bill) bending in hummingbirds have been considered an optical illusion, yet a recent description of out-of-phase opening and closing between their bill base and tip suggests a genuine capacity for bill bending. We investigate bill kinematics during nectar feeding in six species of hummingbirds. We employed geometric morphometrics to identify bending zones and combined these data with measurements of bill flexural rigidity from micro-computed tomography scans to better understand the flexing mechanism. We found that the mandible remains in place throughout the licking cycle, while the maxilla undergoes significant shape deformation, such that the distal portion of the upper bill bends upwards. We propose that bill bending is a key component of the drinking mechanism in hummingbirds, allowing the coordination of bill function (distal wringing and basal expansion) and tongue function (raking/squeegeeing) during intra-oral transport. We present a fluid analysis that reveals a combination of pressure-driven (Poiseuille) and boundary-driven (Couette) flows, which have previously been thought to represent alternative drinking mechanisms. Bill bending allows for separation of the bill tips while maintaining a tightly closed middle section of the bill, enabling nectar exploitation in long and narrow flowers that can exclude less efficient pollinators.

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).)

Terrestrial carnivorans, with their diverse diets and unique adaptations such as the carnassial tooth, offer insights into the connections between functional traits and the climatic and environmental conditions they inhabit. They shed light on functional trait-environment relationships at the highest trophic levels across a broad range of environmental conditions. In this study, we evaluate the relationship between relative blade length (RBL) of the lower carnassial tooth, a key dietary adaptation among terrestrial carnivorans for slicing and grinding food items, and climate. We propose RBL as an ecometric trait and test the hypothesis that community-level RBL is correlated with climate and mediated by environmental effects on food availability. Our findings show that communities with higher mean and broader variance of RBL are typically located in warmer and wetter climates, suggesting a relationship between carnivoran dietary diversity and climate. Conversely, communities with a lower mean and narrower variance of RBL predominantly occupy cooler, drier places. This indicates that community-level carnivoran dietary traits have the potential to serve as indicators of environmental conditions. Given the robust fossil record associated with carnivorans, we also show how RBL can be used as a proxy for reconstructing paleoclimates by examining trait change at seven sites in North America to estimate changes in temperature and precipitation over time in relation to changes in carnivoran community assembly. Understanding the nature of trait-environment relationships can help us anticipate biological impacts of ongoing environmental change and the geographic regions at the greatest risk of ecological disruption., Competing Interests: We have no competing interests to declare., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Squamates have independently evolved an elongate, limb-reduced body form numerous times. This transition has been proposed to involve either changes to regulatory gene expression or downstream modification of target enhancers to produce a homogeneous, deregionalized axial skeleton. Analysis of vertebral morphology has suggested that regionalization is maintained in snake-like body forms, but morphological variation in the other primary component of the axial skeleton, the dorsal ribs, has not been previously examined. We quantified rib morphology along the anterior-posterior axis in limbed and snake-like squamates to test different regionalization models. We find that the relative position of regional boundaries remains consistent across taxa of differing body types, including in the homoplastic evolution of snake-like body forms. The consistent retention of regional boundaries in this primaxial domain is uncorrelated with more plastic abaxial region markers. Rather than loss of regions, rib shape at the anterior and posterior of the axis converges on those in the middle, resulting in axial regions being distinguishable by allometric shape changes rather than by discrete morphologies. This complexity challenges notions of deregionalization, revealing a nuanced evolutionary history shaped by shared functions.

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.

Complex patterns of genome evolution associated with the end-Cretaceous [Cretaceous-Paleogene (K-Pg)] mass extinction limit our understanding of the early evolutionary history of modern birds. Here, we analyzed patterns of avian molecular evolution and identified distinct macroevolutionary regimes across exons, introns, untranslated regions, and mitochondrial genomes. Bird clades originating near the K-Pg boundary exhibited numerous shifts in the mode of molecular evolution, suggesting a burst of genomic heterogeneity at this point in Earth's history. These inferred shifts in substitution patterns were closely related to evolutionary shifts in developmental mode, adult body mass, and patterns of metabolic scaling. Our results suggest that the end-Cretaceous mass extinction triggered integrated patterns of evolution across avian genomes, physiology, and life history near the dawn of the modern bird radiation.

The driving forces behind the evolution of early metazoans are not well understood, but key insights into their ecology and evolution can be gained through ecological analyses of the in situ, sessile communities of the Avalon assemblage in the Ediacaran (~565 million years ago). Community structure in the Avalon is thought to be underpinned by epifaunal tiering and ecological succession, which we investigate in this study in 18 Avalon communities. Here we found that Avalon communities form four distinctive Community Types irrespective of succession processes, which are instead based on the dominance of morphologically distinct taxa, and that tiering is prevalent in three of these Community Types. Our results are consistent with emergent neutrality, whereby ecologically specialized morphologies evolve as a consequence of neutral (stochastic or reproductive) processes within niches, leading to generalization within the frond-dominated Community Type. Our results provide an ecological signature of the first origination and subsequent loss of disparate morphologies, probably as a consequence of community restructuring in response to ecological innovation. This restructuring led to the survival of non-tiered frondose generalists over tiered specialists, even into the youngest Ediacaran assemblages. Such frondose body plans also survive beyond the Ediacaran-Cambrian transition, perhaps due to the greater resilience afforded to them by their alternative ecological strategies., (© 2024. The Author(s).)

Human-induced environmental change has caused widespread loss of species that support important functions for ecosystems and society. For example, vertebrate scavengers contribute to the functional health of ecosystems and provide services to agricultural landscapes by removing carcasses and associated pests. Widespread extirpation of native Australian mammals since the arrival of Europeans in Australia has removed many scavenging species from landscapes, while scavenging mammals such as European red foxes ( Vulpes vulpes ) have been introduced. In much of Australia, squamate reptiles are the largest native terrestrial scavengers remaining, where large native mammals are extinct and conservation management is being undertaken to remove invasive mammals. The contribution of reptiles to scavenging functions is not well understood. In this study, we investigated the ecosystem functions provided by large reptiles as scavengers to better understand how populations can be managed to support ecosystem services. We investigated the ecosystem services provided by vertebrate scavengers in Australian coastal mallee ecosystems, focusing on the heath goanna ( Varanus rosenbergi ), the only extant native terrestrial scavenger in the region. We carried out exclosure experiments, isolating the scavenging activity of different taxonomic groups to quantify the contribution of different taxa to scavenging services, specifically the removal of rat carcasses, and its impact on the occurrence of agriculturally damaging blowflies. We compared areas with different native and invasive scavenger communities to investigate the impact of invasive species removal and native species abundance on scavenging services. Our results indicated that vertebrate scavenging significantly contributes to carcass removal and limitation of necrophagous fly breeding in carcasses and that levels of removal are higher in areas associated with high densities of heath goannas and low densities of invasive mammals. Therefore, augmentation of heath goanna populations represents a promising management strategy to restore and maximize scavenging ecosystem services., Competing Interests: The authors declare no competing interests., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

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.)

Late Ediacaran strata from Newfoundland, Canada (~574-560 Ma) document near-census palaeocommunities of some of the earliest metazoans. Such preservation enables reproductive strategies to be inferred from the spatial distribution of populations of fossilized benthic organisms, previously revealing the existence of both propagule and stoloniferous reproductive modes among Ediacaran frondose taxa. Here, we describe 'conga lines': linear arrangements of more than three closely spaced fossil specimens. We calculate probabilistic models of point maps of 13 fossil-bearing bedding surfaces and show that four surfaces contain conga lines that are not the result of chance alignments. We then test whether these features could result from passive pelagic propagules settling in the lee of an existing frond, using computational fluid dynamics and discrete phase modelling. Under Ediacaran palaeoenvironmental conditions, preferential leeside settlement at the spatial scale of the conga lines is unlikely. We therefore conclude that these features are novel and do not reflect previously described reproductive strategies employed by Ediacaran organisms, suggesting the use of mixed reproductive strategies in the earliest animals. Such strategies enabled Ediacaran frondose taxa to act as reproductive generalists and may be an important facet of early metazoan evolution., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

In birds, the columella is the only bony element of the sound conducting apparatus, conveying vibrations of the cartilaginous extracolumella to the fluid of the inner ear. Although avian columellar morphology has attracted some attention over the past century, it nonetheless remains poorly described in the literature. The few existing studies mostly focus on morphological descriptions in relatively few taxa, with no taxonomically broad surveys yet published. Here we use observations of columellae from 401 extant bird species to provide a comprehensive survey of columellar morphology in a phylogenetic context. We describe the columellae of several taxa for the first time and identify derived morphologies characterizing higher-level clades based on current phylogenies. In particular, we identify a derived columellar morphology diagnosing a major subclade of Accipitridae. Within Suliformes, we find that Fregatidae, Sulidae, and Phalacrocoracidae share a derived morphology that is absent in Anhingidae, suggesting a secondary reversal. Phylogenetically informed comparisons allow recognition of instances of homoplasy, including the distinctive bulbous columellae in suboscine passerines and taxa belonging to Eucavitaves, and bulging footplates that appear to have evolved at least twice independently in Strigiformes. We consider phylogenetic and functional factors influencing avian columellar morphology, finding that aquatic birds possess small footplates relative to columellar length, possibly related to hearing function in aquatic habitats. By contrast, the functional significance of the distinctive bulbous basal ends of the columellae of certain arboreal landbird taxa remains elusive., (© 2023 American Association for Anatomy.)

Warmer temperatures and higher sea level than today characterized the Last Interglacial interval [Pleistocene, 128 to 116 thousand years ago (ka)]. This period is a remarkable deep-time analog for temperature and sea-level conditions as projected for 2100 AD, yet there has been no evidence of fossil assemblages in the equatorial Atlantic. Here, we report foraminifer, metazoan (mollusks, bony fish, bryozoans, decapods, and sharks among others), and plant communities of coastal tropical marine and mangrove affinities, dating precisely from a ca. 130 to 115 ka time interval near the Equator, at Kourou, in French Guiana. These communities include ca. 230 recent species, some being endangered today and/or first recorded as fossils. The hyperdiverse Kourou mollusk assemblage suggests stronger affinities between Guianese and Caribbean coastal waters by the Last Interglacial than today, questioning the structuring role of the Amazon Plume on tropical Western Atlantic communities at the time. Grassland-dominated pollen, phytoliths, and charcoals from younger deposits in the same sections attest to a marine retreat and dryer conditions during the onset of the last glacial (ca. 110 to 50 ka), with a savanna-dominated landscape and episodes of fire. Charcoals from the last millennia suggest human presence in a mosaic of modern-like continental habitats. Our results provide key information about the ecology and biogeography of pristine Pleistocene tropical coastal ecosystems, especially relevant regarding the-widely anthropogenic-ongoing global warming., Competing Interests: Competing interests statement:The authors declare no competing interest.

In birds, the quadrate connects the mandible and skull, and plays an important role in cranial kinesis. Avian quadrate morphology may therefore be assumed to have been influenced by selective pressures related to feeding ecology, yet large-scale variation in quadrate morphology and its potential relationship with ecology have never been quantitatively investigated. Here, we used geometric morphometrics and phylogenetic comparative methods to quantify morphological variation of the quadrate and its relationship with key ecological features across a wide phylogenetic sample. We found non-significant associations between quadrate shape and feeding ecology across different scales of phylogenetic comparison; indeed, allometry and phylogeny exhibit stronger relationships with quadrate shape than ecological features. We show that similar quadrate shapes are associated with widely varying dietary ecologies (one-to-many mapping), while divergent quadrate shapes are associated with similar dietary ecologies (many-to-one mapping). Moreover, we show that the avian quadrate evolves as an integrated unit and exhibits strong associations with the morphologies of neighbouring bones. Our results collectively illustrate that quadrate shape has evolved jointly with other elements of the avian kinetic system, with the major crown bird lineages exploring alternative quadrate morphologies, highlighting the potential diagnostic value of quadrate morphology in investigations of bird systematics.

The origin of crown birds (Neornithes) remains contentious owing to conflicting divergence time hypotheses obtained from alternative sources of data. The fossil record suggests limited diversification of Neornithes in the Late Mesozoic and a substantial radiation in the aftermath of the Cretaceous-Palaeogene (K-Pg) mass extinction, approximately 66 Ma. Molecular clock studies, however, have yielded estimates for neornithine origins ranging from the Early Cretaceous (130 Ma) to less than 10 Myr before the K-Pg. We use Bayes factors to compare the fit of node ages from different molecular clock studies to an independent morphological dataset. Our results allow us to reject scenarios of crown bird origins deep in the Early Cretaceous, as well as an origin of crown birds within the last 10 Myr of the Cretaceous. The scenario best supported by our analyses is one where Neornithes originated between the Early and Late Cretaceous ( ca 100 Ma), while numerous divergences within major neoavian clades either span or postdate the K-Pg. This study affirms the importance of the K-Pg on the diversification of modern birds, and the potential of combined-evidence tip-dating analyses to illuminate recalcitrant 'rocks versus clocks' debates.

Palaeognathae, the extant avian clade comprising the flightless ratites and flight-capable tinamous (Tinamidae), is the sister group to all other living birds, and recent phylogenetic studies illustrate that tinamous are phylogenetically nested within a paraphyletic assemblage of ratites. As the only extant palaeognaths that have retained the ability to fly, tinamous may provide key information on the nature of the flight apparatus of ancestral crown palaeognaths-and, in turn, crown birds-as well as insight into convergent modifications to the wing apparatus among extant ratite lineages. To reveal new information about the musculoskeletal anatomy of tinamous and facilitate development of computational biomechanical models of tinamou wing function, we generated a three-dimensional musculoskeletal model of the flight apparatus of the extant Andean tinamou (Nothoprocta pentlandii) using diffusible iodine-based contrast-enhanced computed tomography (diceCT). Origins and insertions of the pectoral flight musculature of N. pentlandii are generally consistent with those of other extant volant birds specialized for burst flight, and the entire suite of presumed ancestral neornithine flight muscles are present in N. pentlandii with the exception of the biceps slip. The pectoralis and supracoracoideus muscles are robust, similar to the condition in other extant burst-flying birds such as many extant Galliformes. Contrary to the condition in most extant Neognathae (the sister clade to Palaeognathae), the insertion of the pronator superficialis has a greater distal extent than the pronator profundus, although most other anatomical observations are broadly consistent with the conditions observed in extant neognaths. This work will help form a basis for future comparative studies of the avian musculoskeletal system, with implications for reconstructing the flight apparatus of ancestral crown birds and clarifying musculoskeletal modifications underlying the convergent origins of ratite flightlessness., (© 2023 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

The first arrival dates of 31 species of migrant birds in the Tatarstan Republic of Russia were monitored for the 34-year period from 1989-2022. Trends in first arrival date were evaluated using regression against the year value. Patterns in arrival data with respect to species traits (habitat, migration distance, body weight, etc.) were evaluated using redundancy analysis. Relationships between first arrival dates and Tatarstan temperatures were also evaluated using regression methods of first-arrival date on monthly mean temperatures. Almost all (28 of 31) species revealed a significantly earlier migration arrival date; however, associations between arrival patterns and species traits were equivocal. Warmer temperatures were significantly associated with earlier arrival in 26 of the 31 species, but the relationship was insufficient to explain the average 11-day advance in species. For these species and in this location only the timing and location of arrival are well recorded; the exact wintering areas and migration routes, and the timing of these phases are less well understood. When these become better known, an investigation of the influence of environmental conditions (including temperature) on departure timing and passage timing and speed is recommended.

Mammalian megafauna have been critical to the functioning of Earth's biosphere for millions of years. However, since the Plio-Pleistocene, their biodiversity has declined concurrently with dramatic environmental change and hominin evolution. While these biodiversity declines are well-documented, their implications for the ecological function of megafaunal communities remain uncertain. Here, we adapt ecometric methods to evaluate whether the functional link between communities of herbivorous, eastern African megafauna and their environments (i.e., functional trait-environment relationships) was disrupted as biodiversity losses occurred over the past 7.4 Ma. Herbivore taxonomic and functional diversity began to decline during the Pliocene as open grassland habitats emerged, persisted, and expanded. In the mid-Pleistocene, grassland expansion intensified, and climates became more variable and arid. It was then that phylogenetic diversity declined, and the trait-environment relationships of herbivore communities shifted significantly. Our results divulge the varying implications of different losses in megafaunal biodiversity. Only the losses that occurred since the mid-Pleistocene were coincident with a disturbance to community ecological function. Prior diversity losses, conversely, occurred as the megafaunal species and trait pool narrowed towards those adapted to grassland environments., (© 2023. The Author(s).)

In birds and other reptiles, the quadrate acts as a hinge between the lower jaw and the skull and plays an important role in avian cranial kinesis. Though previous studies have qualitatively described substantial variation in quadrate morphology among birds, none have attempted to quantify evolutionary changes in quadrate shape. Here, we investigate geometric evolution of the quadrate in Galloanserae, a major clade of extant birds uniting chicken-like (Galliformes) and duck-like (Anseriformes) fowl. We quantified morphological variation in the quadrate across 50 extant galloanseran species covering all major extant subclades using three-dimensional geometric morphometrics, and performed ancestral shape reconstructions in the context of an up-to-date neornithine phylogeny. We find that our results based only on extant quadrates may overlook plesiomorphic features captured by fossil taxa, resulting in an ancestral state reconstruction for Galloanserae that is seemingly an approximation of the average shape of the extant data set. By contrast, analyses incorporating early fossil galloanseran quadrates (from taxa such as Asteriornis, Presbyornis, and Conflicto) result in ancestral geometric reconstructions more similar to the morphology of extant galliforms, indicating that the quadrate of the last common ancestor of galloanserans may have been more morphologically and functionally similar to those of extant galliforms than to extant anseriforms. These results generally corroborate previous inferences of galloanseran quadrate plesiomorphies and identify several additional plesiomorphic features of the galloanseran quadrate for the first time. Our results illustrate the importance of incorporating fossil taxa into ancestral shape reconstructions and help elucidate important aspects of the morphology and function of the avian feeding apparatus early in crown bird evolutionary history., (© 2023 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

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.

Avian skeletal morphology is associated with locomotor function, including flight style, swimming and terrestrial locomotion, and permits informed inferences on locomotion in extinct taxa. The fossil taxon Ichthyornis (Avialae: Ornithurae) has long been regarded as highly aerial, with flight similar to terns or gulls (Laridae), and skeletal features resembling foot-propelled diving adaptations. However, rigorous testing of locomotor hypotheses has yet to be performed on Ichthyornis , despite its notable phylogenetic position as one of the most crownward stem birds. We analysed separate datasets of three-dimensional sternal shape (geometric morphometrics) and skeletal proportions (linear measurements across the skeleton), to examine how well these data types predict locomotor traits in Neornithes. We then used this information to infer locomotor capabilities of Ichthyornis. We find strong support for both soaring and foot-propelled swimming capabilities in Ichthyornis. Further, sternal shape and skeletal proportions provide complementary information on avian locomotion: skeletal proportions allow better predictions of the capacity for flight, whereas sternal shape predicts variation in more specific locomotor abilities such as soaring, foot-propelled swimming and escape burst flight. These results have important implications for future studies of extinct avialan ecology and underscore the importance of closely considering sternum morphology in investigations of fossil bird locomotion.

The hyper-diverse clade Passeriformes (crown group passerines) comprises over half of extant bird diversity, yet disproportionately few studies have targeted passerine comparative anatomy on a broad phylogenetic scale. This general lack of research attention hinders efforts to interpret the passerine fossil record and obscures patterns of morphological evolution across one of the most diverse clades of extant vertebrates. Numerous potentially important crown passeriform fossils have proven challenging to place phylogenetically, due in part to a paucity of phylogenetically informative characters from across the passerine skeleton. Here, we present a detailed analysis of the morphology of extant passerine carpometacarpi, which are relatively abundant components of the passerine fossil record. We sampled >70% of extant family-level passerine clades (132 extant species) as well as several fossils from the Oligocene of Europe and scored them for 54 phylogenetically informative carpometacarpus characters optimised on a recently published phylogenomic scaffold. We document a considerable amount of previously undescribed morphological variation among passerine carpometacarpi, and, despite high levels of homoplasy, our results support the presence of representatives of both crown Passeri and crown Tyranni in Europe during the Oligocene., (© 2022 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Understanding the relationships between functional traits and environment is increasingly important for assessing ecosystem health and forecasting biotic responses to future environmental change. Taxon-free analyses of functional traits (ecometrics) allow for testing the performance of such traits through time, utilizing both the fossil record and paleoenvironmental proxies. Here, we test the role of body size as a functional trait with respect to climate, using turtles as a model system. We examine the influence of mass-specific metabolic rate as a functional factor in the sorting of body size with environmental temperature and investigate the utility of community body size composition as an ecometric correlated to climate variables. We then apply our results to the fossil record of the Plio-Pleistocene Shungura Formation in Ethiopia. Results show that turtle body sizes scale with mass-specific metabolic rate for larger taxa, but not for the majority of species, indicating that metabolism is not a primary driver of size. Body size ecometrics have stronger predictive power at continental than at global scales, but without a single, dominant predictive functional relationship. Application of ecometrics to the Shungura fossil record suggests that turtle paleocommunity ecometrics coarsely track independent paleoclimate estimates at local scales. We hypothesize that both human disruption and biotic interactions limit the ecometric fit of size to climate in this clade. Nonetheless, examination of the consistency of trait-environment relationships through deep and shallow time provides a means for testing anthropogenic influences on ecosystems.

Rangeomorphs are among the oldest putative eumetazoans known from the fossil record. Establishing how they fed is thus key to understanding the structure and function of the earliest animal ecosystems. Here, we use computational fluid dynamics to test hypothesized feeding modes for the fence-like rangeomorph Pectinifrons abyssalis , comparing this to the morphologically similar extant carnivorous sponge Chondrocladia lyra . Our results reveal complex patterns of flow around P. abyssalis unlike those previously reconstructed for any other Ediacaran taxon. Comparisons with C . lyra reveal substantial differences between the two organisms, suggesting they converged on a similar fence-like morphology for different functions. We argue that the flow patterns recovered for P. abyssalis do not support either a suspension feeding or osmotrophic feeding habit. Instead, our results indicate that rangeomorph fronds may represent organs adapted for gas exchange. If correct, this interpretation could require a dramatic reinterpretation of the oldest macroscopic animals., Competing Interests: The authors declare no competing interests., (© 2023 The Author(s).)

The bony palate diagnoses the two deepest clades of extant birds: Neognathae and Palaeognathae 1-5 . Neognaths exhibit unfused palate bones and generally kinetic skulls, whereas palaeognaths possess comparatively rigid skulls with the pterygoid and palatine fused into a single element, a condition long considered ancestral for crown birds (Neornithes) 3,5-8 . However, fossil evidence of palatal remains from taxa close to the origin of Neornithes is scarce, hindering strong inferences regarding the ancestral condition of the neornithine palate. Here we report a new taxon of toothed Late Cretaceous ornithurine bearing a pterygoid that is remarkably similar to those of the extant neognath clade Galloanserae (waterfowl + landfowl). Janavis finalidens, gen. et sp. nov., is generally similar to the well-known Mesozoic ornithurine Ichthyornis in its overall morphology, although Janavis is much larger and exhibits a substantially greater degree of postcranial pneumaticity. We recovered Janavis as the first-known well-represented member of Ichthyornithes other than Ichthyornis, clearly substantiating the persistence of the clade into the latest Cretaceous 9 . Janavis confirms the presence of an anatomically neognathous palate in at least some Mesozoic non-crown ornithurines 10-12 , suggesting that pterygoids similar to those of extant Galloanserae may be plesiomorphic for crown birds. Our results, combined with recent evidence on the ichthyornithine palatine 12 , overturn longstanding assumptions about the ancestral crown bird palate, and should prompt reevaluation of the purported galloanseran affinities of several bizarre early Cenozoic groups such as the 'pseudotoothed birds' (Pelagornithidae) 13-15 ., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Among terrestrial vertebrates, only crown birds (Neornithes) rival mammals in terms of relative brain size and behavioural complexity. Relatedly, the anatomy of the avian central nervous system and associated sensory structures, such as the vestibular system of the inner ear, are highly modified with respect to those of other extant reptile lineages. However, a dearth of three-dimensional Mesozoic fossils has limited our knowledge of the origins of the distinctive endocranial structures of crown birds. Traits such as an expanded, flexed brain, a ventral connection between the brain and spinal column, and a modified vestibular system have been regarded as exclusive to Neornithes. Here, we demonstrate all of these 'advanced' traits in an undistorted braincase from an Upper Cretaceous enantiornithine bonebed in southeastern Brazil. Our discovery suggests that these crown bird-like endocranial traits may have originated prior to the split between Enantiornithes and the more crownward portion of avian phylogeny over 140 Ma, while coexisting with a remarkably plesiomorphic cranial base and posterior palate region. Altogether, our results support the interpretation that the distinctive endocranial morphologies of crown birds and their Mesozoic relatives are affected by complex trade-offs between spatial constraints during development.

Carcasses of common pheasants (Phasianus colchicus) killed by hunters using shotguns are widely used or sold in the United Kingdom and elsewhere for human consumption. Almost all of the birds are shot using shotgun pellets composed principally of lead (Pb). Lead shotgun pellets often fragment on impact within the bodies of gamebirds, leaving small lead particles in the meat that are difficult for consumers to detect and remove and from which a greater proportion of lead is likely to be absorbed. Chronic exposure to even low levels of lead is associated with negative health effects in humans and especially in groups particularly vulnerable to the effects of lead, which include young children and pregnant women. Our study used a high-resolution computerised tomography X-ray scanner to locate, in three dimensions, metal fragments embedded within carcasses of eight pheasants sold for human consumption in the UK. Small radio-dense fragments (<2 mm diameter), assumed to be metallic lead, were present in all of the pheasant carcasses examined (mean number: 39 per carcass) and many were too small (<0.1 mm diameter) and too distant from the nearest large shotgun pellet for it to be practical for consumers to detect and remove them without discarding a large proportion of otherwise usable meat. Consumers of carcasses of pheasants killed using lead shotgun ammunition are likely to be exposed to elevated levels of dietary lead, even if careful food preparation is practiced to remove shotgun pellets and the most damaged tissue., Competing Interests: The authors have declared that no competing interests exist.

Global temperatures are rising at an unprecedented rate, but environmental responses are often difficult to recognize and quantify. Long-term observations of plant phenology, the annually recurring sequence of plant developmental stages, can provide sensitive measures of climate change and important information for ecosystem services. Here, we present 419 354 recordings of the first flowering date from 406 plant species in the UK between 1753 and 2019 CE. Community-wide first flowering advanced by almost one month on average when comparing all observations before and after 1986 ( p < 0.0001). The mean first flowering time is 6 days earlier in southern than northern sites, 5 days earlier under urban than rural settings, and 1 day earlier at lower than higher elevations. Compared to trees and shrubs, the largest lifeform-specific phenological shift of 32 days is found in herbs, which are generally characterized by fast turnover rates and potentially high levels of genetic adaptation. Correlated with January-April maximum temperatures at -0.81 from 1952-2019 ( p < 0.0001), the observed trends (5.4 days per decade) and extremes (66 days between the earliest and latest annual mean) in the UK's first flowering dataset can affect the functioning and productivity of ecosystems and agriculture.

The axial skeleton of tetrapods is organized into distinct anteroposterior regions of the vertebral column (cervical, trunk, sacral, and caudal), and transitions between these regions are determined by colinear anterior expression boundaries of Hox5/6 , -9 , -10 , and -11 paralogy group genes within embryonic paraxial mesoderm. Fishes, conversely, exhibit little in the way of discrete axial regionalization, and this has led to scenarios of an origin of Hox -mediated axial skeletal complexity with the evolutionary transition to land in tetrapods. Here, combining geometric morphometric analysis of vertebral column morphology with cell lineage tracing of hox gene expression boundaries in developing embryos, we recover evidence of at least five distinct regions in the vertebral skeleton of a cartilaginous fish, the little skate ( Leucoraja erinacea ). We find that skate embryos exhibit tetrapod-like anteroposterior nesting of hox gene expression in their paraxial mesoderm, and we show that anterior expression boundaries of hox5/6 , hox9 , hox10 , and hox11 paralogy group genes predict regional transitions in the differentiated skate axial skeleton. Our findings suggest that hox- based axial skeletal regionalization did not originate with tetrapods but rather has a much deeper evolutionary history than was previously appreciated., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

The altricial-precocial spectrum describes patterns of variation in avian developmental mode that greatly influence avian life histories. Appraising a given species' position on this spectrum is therefore fundamental to understanding patterns of avian life history evolution. However, evaluating avian developmental mode remains a relatively subjective task reliant on untested assumptions, including the notion that developmental strategies are distributed along a single dimension of statistical variation. Here, we present a quantitative multivariate framework that objectively discriminates among meaningfully different modes of avian development. We gathered information on seven hatchling and post-hatching traits for up to 4000 extant bird species, and find that most traits related to developmental mode show high phylogenetic signal and little intraclade variation, allowing unknown values to be reliably interpolated. Principal component analyses (PCAs) of these traits illustrate that most variation in hatchling state can be quantified along one dimension of trait space. However, our PCAs also reveal an important second dimension explaining variation in post-hatching behavior, enabling factors related to hatchling state and post-hatching behavior to be disentangled. In order to facilitate future macroevolutionary studies of variation in avian developmental strategies, as well as explorations of covariation between developmental mode and other aspects of avian biology, we present PC scores for 9993 extant avian species., (© 2021 The Authors. Evolution © 2021 The Society for the Study of Evolution.)

The Cretaceous-Paleogene (K-Pg) mass extinction 66 million years ago was characterized by a worldwide ecological catastrophe and rapid species turnover. Large-scale devastation of forested environments resulting from the Chicxulub asteroid impact likely influenced the evolutionary trajectories of multiple clades in terrestrial environments, and it has been hypothesized to have biased survivorship in favour of nonarboreal lineages across the K-Pg boundary. Here, we evaluate patterns of substrate preferences across the K-Pg boundary among crown group mammals, a group that underwent rapid diversification following the mass extinction. Using Bayesian, likelihood, and parsimony reconstructions, we identify patterns of mammalian ecological selectivity that are broadly similar to those previously hypothesized for birds. Models based on extant taxa indicate predominant K-Pg survivorship among semi- or nonarboreal taxa, followed by numerous independent transitions to arboreality in the early Cenozoic. However, contrary to the predominant signal, some or all members of total-clade Euarchonta (Primates + Dermoptera + Scandentia) appear to have maintained arboreal habits across the K-Pg boundary, suggesting ecological flexibility during an interval of global habitat instability. We further observe a pronounced shift in character state transitions away from plesiomorphic arboreality associated with the K-Pg transition. Our findings are consistent with the hypothesis that predominantly nonarboreal taxa preferentially survived the end-Cretaceous mass extinction, and emphasize the pivotal influence of the K-Pg transition in shaping the early evolutionary trajectories of extant terrestrial vertebrates., Competing Interests: The authors declare no conflict of interest., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Natural succession of vegetation on abandoned farmland provides opportunities for passive rewilding to re-establish native woodlands, but in Western Europe the patterns and outcomes of vegetation colonisation are poorly known. We combine time series of field surveys and remote sensing (lidar and photogrammetry) to study woodland development on two farmland fields in England over 24 and 59 years respectively: the New Wilderness (2.1 ha) abandoned in 1996, and the Old Wilderness (3.9 ha) abandoned in 1961, both adjacent to ancient woodland. Woody vegetation colonisation of the New Wilderness was rapid, with 86% vegetation cover averaging 2.9 m tall after 23 years post-abandonment. The Old Wilderness had 100% woody cover averaging 13.1 m tall after 53 years, with an overstorey tree-canopy (≥ 8 m tall) covering 91%. By this stage, the structural characteristics of the Old Wilderness were approaching those of neighbouring ancient woodlands. The woody species composition of both Wildernesses differed from ancient woodland, being dominated by animal-dispersed pedunculate oak Quercus robur and berry-bearing shrubs. Tree colonisation was spatially clustered, with wind-dispersed common ash Fraxinus excelsior mostly occurring near seed sources in adjacent woodland and hedgerows, and clusters of oaks probably resulting from acorn hoarding by birds and rodents. After 24 years the density of live trees in the New Wilderness was 132/ha (57% oak), with 390/ha (52% oak) in the Old Wilderness after 59 years; deadwood accounted for 8% of tree stems in the former and 14% in the latter. Passive rewilding of these 'Wilderness' sites shows that closed-canopy woodland readily re-established on abandoned farmland close to existing woodland, it was resilient to the presence of herbivores and variable weather, and approached the height structure of older woods within approximately 50 years. This study provides valuable long-term reference data in temperate Europe, helping to inform predictions of the potential outcomes of widespread abandonment of agricultural land in this region., Competing Interests: The authors have declared that no competing interests exist.

This study provides the summary of the reports of the geographical distribution in the Maritime Antarctic and sub-Antarctic regions of Parochlus steinenii (Gercke, 1889) (Diptera, Chironomidae), the only flying insect occurring naturally in the Antarctic continent. The distribution encompasses the South Shetland Islands (Maritime Antarctic), South Georgia (sub-Antarctic), and parts of the Cape Horn Biosphere Reserve (CHBR, southern Chile). In total 78 occurrence records were identified, 53 from our own records, 19 from the literature, and six from other data present in GBIF. Of the 78 records, 66 are from the South Shetland Islands, eight are from South Georgia, and four from the CHBR. This database was developed as one of the main objectives of two Chilean-funded research projects addressing understanding the effects of climate change on sub-Antarctic and Antarctic insects. It provides dataset documenting the distribution of Parochlus steinenii in the Maritime Antarctic, the sub-Antarctic, and the CHBR in southern South America (Chile). The complete dataset is available in Darwin Core Archive format via the Global Biodiversity Information Facility (GBIF)., (Melisa Gañan, Tamara Contador, Javier Rendoll, Felipe Simoes, Carolina Pérez, Gillian Graham, Simón Castillo, James Kennedy, Peter Convey.)

The acquisition of terrestrial, limb-based locomotion during tetrapod evolution has remained a subject of debate for more than a century 1,2 . Our current understanding of the locomotor transition from water to land is largely based on a few exemplar fossils such as Tiktaalik 3 , Acanthostega 4 , Ichthyostega 5 and Pederpes 6 . However, isolated bony elements may reveal hidden functional diversity, providing a more comprehensive evolutionary perspective 7 . Here we analyse 40 three-dimensionally preserved humeri from extinct tetrapodomorphs that span the fin-to-limb transition and use functionally informed ecological adaptive landscapes 8-10 to reconstruct the evolution of terrestrial locomotion. We show that evolutionary changes in the shape of the humerus are driven by ecology and phylogeny and are associated with functional trade-offs related to locomotor performance. Two divergent adaptive landscapes are recovered for aquatic fishes and terrestrial crown tetrapods, each of which is defined by a different combination of functional specializations. Humeri of stem tetrapods share a unique suite of functional adaptations, but do not conform to their own predicted adaptive peak. Instead, humeri of stem tetrapods fall at the base of the crown tetrapod landscape, indicating that the capacity for terrestrial locomotion occurred with the origin of limbs. Our results suggest that stem tetrapods may have used transitional gaits 5,11 during the initial stages of land exploration, stabilized by the opposing selective pressures of their amphibious habits. Effective limb-based locomotion did not arise until loss of the ancestral 'L-shaped' humerus in the crown group, setting the stage for the diversification of terrestrial tetrapods and the establishment of modern ecological niches 12,13 .

Temperatures approaching or exceeding 20°C have been measured during summer in polar regions at the surfaces of barren fellfield soils under cloudless skies around solar noon. However, despite the upper temperature limit for the growth of cold-adapted microbes-which are abundant in polar soils and have pivotal roles in nutrient cycling-typically being close to this temperature, previous studies have not addressed the consequences of climate change for the metabolism of these organisms in the natural environment. Here in a 5-year field experiment on Alexander Island in the southern maritime Antarctic, we show that the abundance of Pseudogymnoascus roseus, the most widespread decomposer fungus in maritime Antarctic fellfield soils, is reduced by 1-2 orders of magnitude when irrigated and nutrient-amended soils are warmed to >20°C during summer. Laboratory experiments under conditions mimicking those during midsummer in the natural environment indicated that the hyphal extension rates of P. roseus isolates and the activities of five extracellular enzymes are reduced by 54%-96% at high water availability after exposure to temperatures cycling daily from 2 to 21°C and 2 to 24°C, relative to temperatures cycling from 2 to 18°C. Given that the temperatures of surface soils at the study site already reach 19°C during midsummer, the observations reported here suggest that, at predicted rates of warming arising from moderate greenhouse gas emissions, inhibitory effects of climate change on the metabolism of P. roseus could manifest themselves within the next few decades. Furthermore, with peak temperatures at the surfaces of fellfield soils at other maritime Antarctic locations and in High Arctic and alpine regions already exceeding 20°C during summer, the observations suggest that climate warming has the potential to inhibit the growth of other cold-adapted microbes, with negative effects on soils as the Earth's climate continues to warm., (© 2020 The Authors. Global Change Biology published by John Wiley && Sons Ltd.)

A paper published in Global Change Biology in 2006 revealed that phenological responses in 1971-2000 matched the warming pattern in Europe, but a lack of chilling and adaptation in farming may have reversed these findings. Therefore, for 1951-2018 in a corresponding data set, we determined changes as linear trends and analysed their variation by plant traits/groups, across season and time as well as their attribution to warming following IPCC methodology. Although spring and summer phases in wild plants advanced less (maximum advances in 1978-2007), more (~90%) and more significant (~60%) negative trends were present, being stronger in early spring, at higher elevations, but smaller for nonwoody insect-pollinated species. These trends were strongly attributable to winter and spring warming. Findings for crop spring phases were similar, but were less pronounced. There were clearer and attributable signs for a delayed senescence in response to winter and spring warming. These changes resulted in a longer growing season, but a constant generative period in wild plants and a shortened one in agricultural crops. Phenology determined by farmers' decisions differed noticeably from the purely climatic driven phases with smaller percentages of advancing (~75%) trends, but farmers' spring activities were the only group with reinforced advancement, suggesting adaptation. Trends in farmers' spring and summer activities were very likely/likely associated with the warming pattern. In contrast, the advance in autumn farming phases was significantly associated with below average summer warming. Thus, under ongoing climate change with decreased chilling the advancing phenology in spring and summer is still attributable to warming; even the farmers' activities in these seasons mirror, to a lesser extent, the warming. Our findings point to adaptation to climate change in agriculture and reveal diverse implications for terrestrial ecosystems; the strong attribution supports the necessary mediation of warming impacts to the general public., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

The squamate fossil record from Kanapoi reveals generic to higher-order similarities with modern East African herpetofaunas. The record is derived from surface collection and screen washing, and consists primarily of isolated vertebrae with a few maxillary and mandibular elements. The most abundant remains are vertebrae of large-bodied Python that are morphologically similar to extant Python sebae, and vertebrae of Varanus cf. (Varanus niloticus + Varanus exanthematicus). Additional cranial and vertebral remains indicate the presence of lygosomine skinks, indeterminate Varanus, Viperidae, cf. Atractaspididae, and multiple colubrine morphotypes in the Kanapoi ecosystem. Despite similarities with modern herpetofaunas, the Kanapoi record lacks taxa common to other East African records, including agamids, chamaeleonids, amphisbaenians, the elapid Naja, and typhlopids. The overall composition of the Kanapoi squamate record is consistent with paleoenvironments similar to modern shrub savanna habitats. There are no indicators of canopied forest environments in squamate faunal composition. The fossil record of Kanapoi suggests that assembly of squamate faunas of modern East Africa was well underway by the late Neogene., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Ethanol is a natural by-product of the fermentation process of fruit sugars and is occasionally consumed by fruit-eating and tree sap drinking birds. Information on this form of alcohol consumption features in the scientific literature. However, as pets or as wild animals living close to humans in urban habitats, birds have increasing possibilities to consume alcohol from beverages, such as beer, wine or spirits. Some observations have been discussed in a light-hearted manner in mass media and social media, but without any generalization of why some bird species drink the beverages intentionally or unintentionally provided by humans. To check which species and in what circumstances birds drink alcohol and how this is evaluated by humans, we reviewed the scientific literature and analysed videos from YouTube. In total we found and analysed 8 scientific papers and 179 YouTube videos, from which we identified at least 55 species (in some cases not all birds were identified to species level), 11 in the scientific literature and 47 in videos. The distribution of these species over the avian phylogenetic tree suggests that the origin of this convergent behaviour is mainly by human influence. The two data sources differed in the species covered. Videos typically presented interactions of birds with human-provided alcoholic beverages, and were dominated by two groups of intelligent birds: parrots and corvids. The popularity of YouTube videos for a particular species was positively correlated with the general popularity of the species as measured by the number of hits (results listed) on Google. Human responses to the videos were generally very positive and we analysed how the responses were influenced by factors derived from viewing the videos. Moreover, YouTube videos also provide information on at least 47 new bird species not previously mentioned as using alcohol, and our results suggest that parrots in particular can be potentially good candidates for future restricted laboratory studies on the effect of ethanol on birds and their relationship with humans., Competing Interests: The authors declare no conflict of interest.

The known diversity of tetrapods of the Devonian period has increased markedly in recent decades, but their fossil record consists mostly of tantalizing fragments 1-15 . The framework for interpreting the morphology and palaeobiology of Devonian tetrapods is dominated by the near complete fossils of Ichthyostega and Acanthostega; the less complete, but partly reconstructable, Ventastega and Tulerpeton have supporting roles 2,4,16-34 . All four of these genera date to the late Famennian age (about 365-359 million years ago)-they are 10 million years younger than the earliest known tetrapod fragments 5,10 , and nearly 30 million years younger than the oldest known tetrapod footprints 35 . Here we describe Parmastega aelidae gen. et sp. nov., a tetrapod from Russia dated to the earliest Famennian age (about 372 million years ago), represented by three-dimensional material that enables the reconstruction of the skull and shoulder girdle. The raised orbits, lateral line canals and weakly ossified postcranial skeleton of P. aelidae suggest a largely aquatic, surface-cruising animal. In Bayesian and parsimony-based phylogenetic analyses, the majority of trees place Parmastega as a sister group to all other tetrapods.