The vast majority of water-walking animals are tiny invertebrates. Most water running vertebrates are larger and specialized for this behavior. As size increases, the role of surface tension decreases and inertial mechanisms dominate. We report how Anolis sagrei, a small, quadrupedal, arboreal lizard with adhesive pads runs on water, using behavioral observations, high-speed video, and decreasing surface tension with a surfactant. We quantified how frequently anoles ran on water, and if this was affected by reduced surface tension. We also compared their running performance and kinematics in the two types of water. Finally, we tested whether certain morphological features affected performance, and what kinematic variables were used to modulate performance in water. We found that decreasing surface tension resulted in a decreased frequency of running on water, but no change in performance, likely due to kinematic compensation. Specifically, anoles ran successfully in 17% of trials and this decreased to 9% with surfactant. Performance, measured as average velocity, Froude number, and the proportion of the body out of the water were unaffected by reduced surface tension. Anoles used faster slaps and shorter undulatory wavelengths when surface tension was reduced. Relative mass and limb characteristics did not affect water running performance. Increased stride and undulatory frequencies were the primary strategies for increasing water running performance. The frequency of behavior may help reconcile past findings, where decreased surface tension decreased water running performance in house geckos but not basilisks.

Convergence is the evolution of similar phenotypes often due to similar selective pressures or constraints limiting evolutionary options. Snake-like morphologies, characterized by elongated bodies and reduced limbs, have evolved repeatedly among vertebrates, including numerous times in squamate reptiles (lizards and snakes). It has been suggested that elongation facilitates locomotion through substrates while limb reduction typically occurs in clade-specific patterns, but this has not been tested. We compared the fit of a series of habitat- and clade-specific models for the evolution of digits, phalanges. and trunk vertebrae in lizards. We found that species inhabiting fossorial and cluttered habitats differed in numbers of vertebrae, digits, and phalanges from species in other habitats. A model with habitat-specific rates fit best for vertebral evolution, with sand swimmers, litter dwellers, and burrowers having higher rates of vertebral evolution than non-fossorial taxa. However, we found digits and phalanges evolved in a clade-specific manner, with higher rates of limb evolution in certain clades. This suggests that limb reduction in snake-like lizards is dictated by clade-specific constraints. In contrast, fossoriality appears to relax functional constraints on vertebral number, facilitating body form diversification. These results suggest that the relaxation of constraints may be an additional mechanism for convergent evolution.

Many-to-many mapping of form-to-function posits that multiple morphological and physiological traits affect the performance of multiple tasks in an organism, and that redundancy and multitasking occur simultaneously to shape the evolution of an organism's phenotype. Many-to-many mapping is expected to be ubiquitous in nature, yet little is known about how it influences the evolution of organismal phenotype. The F-matrix is a powerful tool to study these issues because it describes how multiple traits affect multiple tasks. We undertook a simulation study using the F-matrix to test how the number of traits and the number of tasks affect trait integration and evolvability, as well as the relationships among tasks. We found that as the number of traits and/or tasks increases, the relationships between the tasks and the integration between the traits become weaker, and that the evolvability of the traits increases, all resulting in a system that is freer to evolve. We also found that as the number of traits increases, performance tradeoffs tend to become weaker, but only to a point. Our work shows that it is important to consider not only multiple traits, but also the multitude of tasks that those traits carry out when studying form-function relationships. We suggest that evolution of these relationships follows functional lines of least resistance, which are less defined in more complex systems, resulting in a mechanism for diversification.

Predator–prey interactions are ubiquitous and under strong selection because of the consequences experienced by both predator and prey if they lose the interaction. Biomechanics and behavior play important roles in the outcome of these interactions, but many studies focus on the prey, restrict the range of behaviors considered, and the role of prey boldness in the outcome is not understood. We used high‐speed video to test for effects of multiple measures of performance and kinematics of both the predator and prey, and boldness of prey on the outcome of interactions between Pike Cichlids ( Crenicichla sp.) and Guppies ( Poecilia reticulata ). We found high variation in the behaviors employed during the predator–prey interactions, including in suction versus raptorial feeding, strike accuracy, and guppy responsiveness. We also found that predators moving relatively slower and prey moving relatively faster were more successful at consuming the prey and evading the predator, respectively. Prey that reacted farther from the predator was more likely to escape predation, but boldness of the prey did not affect the interaction. Our work suggests that a high level of variation in predator–prey interactions is widespread, even when strike and escape behaviors are stereotyped. We also showed that what both the predator and the prey do during an interaction are important in determining the outcome.

Minoritized racial groups in the U.S. have experienced disproportionately higher rates of COVID-19 cases and deaths. Studies have linked structural racism as a critical factor causing these disproportionate health burdens. We analyse the relationships between county-level COVID-19 cases and deaths and five measures of structural racism on Black Americans: Black-White residential segregation, differences in educational attainment, unemployment, incarceration rates, and health insurance coverage between Black and White Americans. When controlling for socioeconomic, demographic, health and behavioural factors significant relationships were found between all measures of structural racism with cases and/or deaths except Black-White differences in health insurance coverage. Black-White disparities in educational attainment and incarceration were the strongest predictors. The results varied greatly across regions of the U.S. We also found strong relationships between COVID-19 and mobility and the proportion of foreign-born non-citizens. This work supports the important need to confront structural racism on multiple fronts to address health disparities.

People of different racial/ethnic backgrounds, demographics, health, and socioeconomic characteristics have experienced disproportionate rates of infection and death due to COVID-19. This study tests if and how county-level rates of infection and death have changed in relation to societal county characteristics through time as the pandemic progressed. This longitudinal study sampled monthly county-level COVID-19 case and death data per 100,000 residents from April 2020 to March 2022, and studied the relationships of these variables with racial/ethnic, demographic, health, and socioeconomic characteristics for 3125 or 97.0% of U.S. counties, accounting for 96.4% of the U.S. population. The association of all county-level characteristics with COVID-19 case and death rates changed significantly through time, and showed different patterns. For example, counties with higher population proportions of Black, Native American, foreign-born non-citizen, elderly residents, households in poverty, or higher income inequality suffered disproportionately higher COVID-19 case and death rates at the beginning of the pandemic, followed by reversed, attenuated or fluctuating patterns, depending on the variable. Patterns for counties with higher White versus Black population proportions showed somewhat inverse patterns. Counties with higher female population proportions initially had lower case rates but higher death rates, and case and death rates become more coupled and fluctuated later in the pandemic. Counties with higher population densities had fluctuating case and death rates, with peaks coinciding with new variants of COVID-19. Counties with a greater proportion of university-educated residents had lower case and death rates throughout the pandemic, although the strength of this relationship fluctuated through time. This research clearly shows that how different segments of society are affected by a pandemic changes through time. Therefore, targeted policies and interventions that change as a pandemic unfolds are necessary to mitigate its disproportionate effects on vulnerable populations, particularly during the first six months of a pandemic.

Although typically possessing four limbs and short bodies, lizards have evolved diverse morphologies, including elongate trunks with tiny limbs. Such forms are hypothesized to aid locomotion in cluttered/fossorial environments but propulsion mechanisms (e.g., the use of body and/or limbs to interact with substrates) and potential body/limb coordination remain unstudied. Here, we use biological experiments, a geometric theory of locomotion, and robophysical models to investigate body-limb coordination in diverse lizards. Locomotor field studies in short-limbed, elongate lizards (Brachymeles and Lerista) and laboratory studies of fully limbed lizards (Uma scoparia and Sceloporus olivaceus) and a snake (Chionactis occipitalis) reveal that body-wave dynamics can be described by a combination of standing and traveling waves; the ratio of the amplitudes of these components is inversely related to the degree of limb reduction and body elongation. The geometric theory (which replaces laborious calculation with diagrams) helps explain our observations, predicting that the advantage of traveling-wave body undulations (compared with a standing wave) emerges when the dominant thrust-generation mechanism arises from the body rather than the limbs and reveals that such soil-dwelling lizards propel via "terrestrial swimming" like sand-swimming lizards and snakes. We test our hypothesis by inducing the use of traveling waves in stereotyped lizards via modulating the ground-penetration resistance. Study of a limbed/undulatory robophysical model demonstrates that a traveling wave is beneficial when propulsion is generated by body-environment interaction. Our models could be valuable in understanding functional constraints on the evolutionary processes of elongation and limb reduction as well as advancing robot designs.

Many animals frequently transition between different media while navigating their heterogeneous environments. These media vary in compliance, moisture content, and other characteristics that affect their physical properties. As a result, animals may need to alter their kinematics to adapt to potential changes in media while maintaining performance during predator escape and foraging. Due to its fluid nature, water is highly compliant, and although usually associated with swimming, water running has evolved in a variety of animals ranging from insects to mammals. While the best studied large water runners are the bipedal basilisk lizards (Basiliscus spp.), other lizards have also been observed to run across the surface of water, namely Hemidactylus platyurus, a house gecko, and in this study, Anolis sagrei, the brown anole. Unlike the basilisk lizard, the primarily arboreal Anolis sagrei is not adapted for water running. Moreover, water running in A. sagrei, similar to that of the house gecko, was primarily quadrupedal. Here, we tested for performance and kinematic differences between aquatic and terrestrial running and if the variance in performance and kinematic variables differed between the two media. We found no difference in average and maximum velocity between running on land and water. We also found that Anolis sagrei had higher hindlimb stride frequencies, decreased duty factor, and shorter stride lengths on water, as well as more erect postures. Finally, we found that most kinematics did not differ in variance between the two media, but of those that were different, almost all were more variable during terrestrial running. Our findings show that animals may be capable of specialized modes of locomotion, even if they are not obviously adapted for them, and that they may do this by modulating their kinematics to facilitate locomotion through novel environments.

Physical principles and laws determine the set of possible organismal phenotypes. Constraints arising from development, the environment, and evolutionary history then yield workable, integrated phenotypes. We propose a theoretical and practical framework that considers the role of changing environments. This 'ecomechanical approach' integrates functional organismal traits with the ecological variables. This approach informs our ability to predict species shifts in survival and distribution and provides critical insights into phenotypic diversity. We outline how to use the ecomechanical paradigm using drag-induced bending in trees as an example. Our approach can be incorporated into existing research and help build interdisciplinary bridges. Finally, we identify key factors needed for mass data collection, analysis, and the dissemination of models relevant to this framework.

Granular substrates ranging from silt to gravel cover much of the Earth's land area, providing an important habitat for fossorial animals. Many of these animals use their heads to penetrate the substrate. Although there is considerable variation in head shape, how head shape affects fossorial locomotor performance in different granular substrates is poorly understood. Here, head shape variation for 152 species of fossorial lizards was quantified for head diameter, slope and pointiness of the snout. The force needed to penetrate different substrates was measured using 28 physical models spanning this evolved variation. Ten substrates were considered, ranging in particle size from 0.025 to 4 mm in diameter and consisting of spherical or angular particles. Head shape evolved in a weakly correlated manner, with snouts that were gently sloped being blunter. There were also significant clade differences in head shape among fossorial lizards. Experiments with physical models showed that as head diameter increased, absolute penetration force increased but force normalized by cross-sectional area decreased. Penetration force decreased for snouts that tapered more gradually and were pointier. Larger and angular particles required higher penetration forces, although intermediate size spherical particles, consistent with coarse sand, required the lowest force. Particle size and head diameter effect were largest, indicating that fossorial burrowers should evolve narrow heads and bodies, and select relatively fine particles. However, variation in evolved head shapes and recorded penetration forces suggests that kinematics of fossorial movement are likely an important factor in explaining evolved diversity.

Evolutionary reversals, including re-evolution of lost structures, are commonly found in phylogenetic studies. However, we lack an understanding of how these reversals happen mechanistically. A snake-like body form has evolved many times in vertebrates, and occasionally a quadrupedal form has re-evolved, including inBrachymeleslizards. We use body form and locomotion data for species ranging from snake-like to quadrupedal to address how a quadrupedal form could re-evolve. We show that large, quadrupedal species are faster at burying and surface locomotion than snake-like species, indicating a lack of expected performance trade-off between these modes of locomotion. Species with limbs use them while burying, suggesting that limbs are useful for burying in wet, packed substrates. Palaeoclimatological data suggest thatBrachymelesoriginally evolved a snake-like form under a drier climate probably with looser soil in which it was easier to dig. The quadrupedal clade evolved as the climate became humid, where limbs and large size facilitated fossorial locomotion in packed soils.

Background Diverse architectures of nervous systems (NSs) such as a plexus in cnidarians or a more centralized nervous system (CNS) in insects and vertebrates are present across Metazoa, but it is unclear what selection pressures drove evolution and diversification of NSs. One underlying aspect of this diversity lies in the cellular and molecular mechanisms driving neurogenesis, i.e. generation of neurons from neural precursor cells (NPCs). In cnidarians, vertebrates, and arthropods, homologs of SoxB and bHLH proneural genes control different steps of neurogenesis, suggesting that some neurogenic mechanisms may be conserved. However, data are lacking for spiralian taxa. Results To that end, we characterized NPCs and their daughters at different stages of neurogenesis in the spiralian annelid Capitella teleta . We assessed cellular division patterns in the neuroectoderm using static and pulse-chase labeling with thymidine analogs (EdU and BrdU), which enabled identification of NPCs that underwent multiple rounds of division. Actively-dividing brain NPCs were found to be apically-localized, whereas actively-dividing NPCs for the ventral nerve cord (VNC) were found apically, basally, and closer to the ventral midline. We used lineage tracing to characterize the changing boundary of the trunk neuroectoderm. Finally, to start to generate a genetic hierarchy, we performed double-fluorescent in-situ hybridization (FISH) and single-FISH plus EdU labeling for neurogenic gene homologs. In the brain and VNC, Ct-soxB1 and Ct-neurogenin were expressed in a large proportion of apically-localized, EdU + NPCs. In contrast, Ct-ash1 was expressed in a small subset of apically-localized, EdU + NPCs and subsurface, EdU − cells, but not in Ct-neuroD + or Ct-elav1 + cells, which also were subsurface. Conclusions Our data suggest a putative genetic hierarchy with Ct-soxB1 and Ct-neurogenin at the top, followed by Ct-ash1 , then Ct-neuroD , and finally Ct-elav1 . Comparison of our data with that from Platynereis dumerilii revealed expression of neurogenin homologs in proliferating NPCs in annelids, which appears different than the expression of vertebrate neurogenin homologs in cells that are exiting the cell cycle. Furthermore, differences between neurogenesis in the head versus trunk of C. teleta suggest that these two tissues may be independent developmental modules, possibly with differing evolutionary trajectories.