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    Area of Habitat maps for the world’s terrestrial birds and mammals

    Knowing the distribution of species is crucial for effective conservation action. However, accurate and high-resolution spatial data are only available for a limited number of species1,2. For mammals and birds, the most comprehensive and widely used global distribution dataset is the set of range maps compiled as part of the assessments for the International Union for Conservation of Nature (IUCN) Red List. These represent each species’ distributional limits and tend to minimize omission errors (i.e. false absences) at the expense of commission errors (i.e. false presences)3,4. Therefore, they often contain sizeable areas not regularly occupied by the species.Maps of the Area of Habitat (AOH; previously known as Extent of Suitable Habitat, ESH) complement range maps by indicating potential occupancy within the range, thereby reducing commission errors5. AOH is defined as ‘the habitat available to a species, that is, habitat within its range’5. These models are produced by subtracting areas unsuitable for the species within their range, using information on each species’ associations with habitat and elevation5,6,7,8. Comprehensive sets of AOH maps have been produced in the past for mammals6 and amphibians7, as well as subsets of birds8,9. The percentage of a species’ range covered by the AOH varies depending on the methodology used to associate species to their habitats, and their habitats to land-cover, the coarseness of the range map, the region in which the species is distributed, and the species’ habitat specialization and elevation limits5. For example, Rondinini et al.6 found that, when considering elevation and land cover features for terrestrial mammals, the AOH comprised, on average, 55% of the range. Ficetola et al.7 obtained a similar percentage when analyzing amphibians (55% for forest species, 42% for open habitat species and 61% for habitat generalists). Beresford et al.8 found that AOH covered a mean of 27.6% of the range maps of 157 threatened African bird species. In 2019, Brooks et al.5 proposed a formal definition and standardized methodology to produce AOH, limiting the inputs to habitat preferences, elevation limits, and geographical range.AOH production requires knowledge of which habitat types a species occurs in and their location within the range1. Information on habitat preference is documented for each species assessed in the IUCN Red List10, following the IUCN Habitats Classification Scheme11. However, the IUCN does not define habitat classes in a spatially explicit way, therefore, we used a recently published translation table that associates IUCN Habitat Classification Scheme classes with land cover classes12. Species’ elevation limits were also extracted from the IUCN Red List.We developed AOH maps for 5,481 terrestrial mammal species and 10,651 terrestrial bird species (Fig. 1). For 1,816 bird species defined by BirdLife International as migratory, we developed separate AOH maps, for the resident, breeding, and non-breeding ranges, according to the migratory distribution of the species (Fig. 2). The maps are presented in a regular latitude/longitude grid with an approximate 100 m resolution at the equator. On average, the AOH covers 66 ± 28% of the geographical range for mammals and 64 ± 27% for birds. We used the resulting AOH maps to produce four global species richness layers for: mammals, birds, globally threatened mammals and globally threatened birds13 (Fig. 3).Fig. 1Spatial distribution maps of Tangara abbas. Maps represent (a) the geographic range21, and (b) the Area of Habitat (AOH) of the species. The AOH was produced by subtracting unsuitable habitats from the geographical range. This species’ habitats are forest and terrestrial artificial habitats and has elevation range of 0 – 1600 m.Full size imageFig. 2Spatial distribution maps of Cardellina rubrifrons, divided into resident, breeding and non-breeding areas for this migratory species. Maps represent (a) the geographic range21, and (b) the Area of Habitat (AOH) of the species. The AOH was produced by subtracting unsuitable habitats from the ranges. This species is a forest species with elevation rangelimits of 1500 – 3100 m.Full size imageFig. 3Global species richness maps for (a) terrestrial mammals (considering 5,481species) and (b) terrestrial birds (considering 10,651 species). Calculated by overlaying all species’ AOH per class, resulting inon the number of species at each grid cell, latitude/longitude grid at a resolution of 1°/1008 or approximately 100 m at the equator (EPSG:4326) with the ellipsoid WGS 1984.Full size imageThe AOH maps presented in this paper are more useful for some purposes than global species distribution models, as they reduce and standardize commissions14. They are especially useful for not well-known and wide-range species. However, we note that for well-known species alternative sources may have more accurate distributions15. Moreover, AOHs are affected by the bias and errors of the underlying data, especially relevant errors associated with documentation of species’ habitats and elevations, and the translation of habitats into land cover classes, given that habitat is a complex multidimensional concept that is challenging to match to land-cover classes12, and that the current version of the IUCN Habitat Classification Scheme on IUCN’s website is described as a draft version11.The AOH maps have multiple conservation applications5,16,17, such as assessing species’ distributions and extinction risk, improving the accuracy of conservation planning, monitoring habitat loss and fragmentation, and guiding conservation actions. AOH has been proposed as an additional spatial metric to be documented in the Red List5, and is used for the identification of Key Biodiversity Areas18. More

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    Analysis of influencing factors of phenanthrene adsorption by different soils in Guanzhong basin based on response surface method

    Surface morphology analysisSEM images were shown in Fig. 1. It showed that the contour of three soils were fairly clear before adsorption. But it became fuzzier and the degree of cementation was increased when phenanthrene was adsorbed on the soils. According to the surface morphology, the silty sand (A) had furrows on the surface before adsorption compared with the fairly smooth without any furrows after adsorption (B). The silts (C) were flaky and the lamellar accumulation decreased (D). The loess (E) had a smooth surface with some flaky and rod like structure, after adsorption (F), the surface of loess increased in clay-like structure.Figure 1SEM micrographs of the three soil samples. (A) Silty sand; (B) Adsorbing 5 h of Silty sand; (C) Silts; (D) Adsorbing 5 h of Silts; (E) Loess; (F) Adsorbing 5 h of Loess.Full size imageAdsorption and desorption experimentsAdsorption and desorption kineticsAdsorption kinetics is one of the most important characteristics governing solute uptake rate and represents adsorption efficiency33. The sorption and desorption kinetics of phenanthrene in three soils were shown in Fig. 2. The results showed that the adsorption processes among all soils were similar. The kinetics of phenanthrene in soils was completed in two steps: a “fast” adsorption and a “slow” adsorption. The adsorption amount increased during 0-18h. It was a rapid reaction from 0 to 200 minutes. From 200 to 600 minutes, the adsorption amount increased slightly into balance. This phenomenon was due to the adsorption of phenanthrene occurred on the surface of soil organic matter. With the increase of time, soil surface adsorption sites were gradually saturated, causing the decrease of adsorption rate until reaching the equilibrium. Phenanthrene was a hydrophobic substance. It was easy to reach the soil surface and adhere to the grain surface. The results were consistent with the study of had also found that the balance time was approximately 18h and the adsorption amount increased with the adsorption reaction time34. Under the same conditions, loess had the highest adsorption capacity, which was mainly due to the highest organic content 18. The maximum phenanthrene sorption capacities ranked as follows: loess > silty sand > silts. As shown in Fig. 2, phenanthrene desorption in soils was relatively quick and its desorption equilibrium time was 3h. To reach an adequate desorption balance while remaining consistent with the adsorption reaction time, the balance time of the adsorption–desorption experiment was set at 18h. Generally, PAHs below 4 cycles could reach the adsorption equilibrium for about 16~24h.Figure 2(a)Adsorption equilibration curves of phenanthrene sorption in soils. (b) Desorption equilibration curves of phenanthrene sorption in soils.Full size imagePseudo-second-order and Elovich models were used to study the phenanthrene adsorption mechanism (Table 3). Phenanthrene sorption kinetics were satisfactorily described by a pseudo-second-order model with coefficients of determination (R2) ranging from 0.99875 to 0.99847, compared with R2 values of 0.26508–0.73901 for the Elovich model. This well-fitting pseudo-second-order model indicated that the rate-limiting step was chemical adsorption, including electronic forces through sharing or exchange of electrons35,36. Moreover, it suggested that sorption was governed by the availability of sorption sites on the soil surfaces instead of by the phenanthrene concentration in solution.Table 3 Constants and coeffients of determination of Pseudo-second-order kinetics and Elovich models of sorption.Full size tableAdsorption and desorption isothermsThe isotherm was used for quantitative analysis of phenanthrene transport from liquid to solid phase and for understanding the nature of interactions between phenanthrene and the soil matrix. The sorption and desorption isotherms of phenanthrene in soils were shown in Fig. 3. The data showed that phenanthrene adsorption and desorption capacities of three soils varied markedly due to their different physicochemical properties. With the increase of phenanthrene concentration, the adsorbed amount increased. At the same temperature, the adsorption capacity of silty sand was minimum while loess was maximum. This is mainly related to the soil physicochemical properties. At the same initial concentration, the temperature increase from 20 °C to 40 °C showed that the adsorption and desorption capacity decreased with temperature increase. On the one hand, the rise of temperature can increase the phenanthrene solubility in the liquid phase. On the other hand, it could reduce various forces between the soil surface and phenanthrene37.Figure 3(a)20 °C adsorption isotherms for phenanthrene in soils. (b)30 °C adsorption isotherms for phenanthrene in soils. (c)40 °C adsorption isotherms for phenanthrene in soils. (d) 20 °C desorption isotherms for phenanthrene in soils. (e) 30 °C desorption isotherms for phenanthrene in soils. (f) 40 °C desorption isotherms for phenanthrene in soils.Full size imageThe Freundlich isotherm was used mainly for adsorption surfaces with nonuniform energy distribution, and the Langmuir isotherm was used for monolayer adsorption on perfectly smooth and homogeneous surfaces38. The experimental data were fitted with the Langmuir and Freundlich adsorption models, and the isotherm parameters logKF, 1/n, KL, qmax and the coefficient of determination (R2) of phenanthrene in soils were listed in Table 4.Table 4 Isotherm parameters for Phenanthrene sorption in soils.Full size tableAs shown in Table 4, according to the coefficients of determination (R2), all soils were better fitted with the Freundlich model, which assumes that phenanthrene sorption and desorption occurs on a heterogeneous surface with the possibility of sorption being multi-layered39. This phenomenon has also been observed in humic acid and nanometer clay mineral40. It showed that the soil adsorption of organic matter was not only surface adsorption but also the process of soil organic matter distribution41,42,43 reached the equilibrium isotherm fitted well with the Freundlich equation when studying the adsorption behavior of aromatic compounds by solids.Adsorption and desorption thermodynamicsTo clarify the adsorption mechanisms, the thermodynamic parameters mentioned earlier were calculated and presented in Table 5. Generally, the value of Gibbs free energy changeΔG0 indicated the spontaneity of a chemical reaction. Therefore, it could evaluate whether sorption was relate to spontaneous interaction44. Negative values of ΔG0 indicated that the feasibility and spontaneous nature. The research was under the temperature range about 293–313 K. For adsorption process, all soils ΔG0 was  0 and desorption ΔH  1, P  temperature  > phenanthrene concentration  > pH. In the interaction, the phenanthrene concentration and organic matter have a significant effect on the silt adsorption rate. The coefficient of determination of the silt complex correlation is R2 = 0.9464, indicating that the response model has a good fit, and the experimental error is within the acceptable range. Adjusting the complex correlation coefficient R2 = 0.8982 indicates that the regression relationship can explain 89.82% of the change in the dependent variable. Therefore, this The model can be used to analyze and predict the effect of different factors on the adsorption rate of phenanthrene.3D response surface analysisIn response surface optimization, the three-dimensional response surface graph reflects the influence of the interaction of the other two variables on the response value, and the slope of the response surface reflects the significance of the interaction of the two variables on the response value. The more significant the interaction effect is on the response value, when the slope is gentle, the effect is not significant. If the contour map is elliptical, it indicates that the interaction between the two variables is significant, and if the contour map is circular, it is not significant46. In addition, the slope and density of the contour line also reflect the influence of the variable on the response value. The steeper the contour line and the greater the density, the greater the influence of the variable on the response value47.

    (1) Loess Fig. 5 is a three-dimensional response surface diagram of the interaction between initial phenanthrene concentration and pH to phenanthrene adsorption on loess. It can be seen from the figure that the slope of the response surface graph is steep, and the contour line is an approximate circle, indicating that the interaction between phenanthrene concentration and pH is not significant for the response value. With the increase of pH, the adsorption rate of phenanthrene on loess showed a slow decline at first to the lowest point at 6, and then gradually increased. When the soil pH was close to 6, with the increase of the initial phenanthrene concentration, the adsorption rate of loess also showed a trend of first decreasing and then increasing. According to the F value, F = 0.337, P = 0.5532  > 0.05, it can be concluded that soil pH and initial phenanthrene concentration of the solution have no significant interaction on the adsorption rate of loess.

    Figure 6 shows the effects of initial phenanthrene concentration and organic matter on phenanthrene adsorption on Loess under the condition that pH value and temperature are at the central point. It can be seen from the figure that the initial phenanthrene concentration and soil organic matter contour are steep, indicating that their interaction is significant. The range of phenanthrene adsorption rate is 70 ~ 95, and the change of surface is steep. From the Loess error analysis, it can be seen that if f value is 6.05 and P value is 0.0275  More

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    Comparison of the effects of litter decomposition process on soil erosion under simulated rainfall

    Study area descriptionYangtze River Basin is situated in central China (Fig. 1). Its geographical coordinates are between 30° 48′ 30″–31° 02′ 30″ N and 112° 48′ 45″–113° 03′ 45″ E. Taizishan is located in the transition zone between the north and south of China, with an altitude of 403–467.4 m. It belongs to the subtropical monsoon humid climate zone and has obvious karst landforms. The farm area is 7576 hectares, the forest coverage rate is 82.0%, and the vegetation is mainly Masson pine, fir, and various broad-leaved tree species. Increased forest coverage reduces sediment production30. The soil is mainly viscous yellow–brown soil and loess parent material. Rain is concentrated in summer, with an average annual rainfall of 1094.6 mm and an average annual temperature of 16.4 °C. Rainfall-related flood risk increased in the Yangtze River Delta in recent years31.The study was based in a Pinus massoniana forest in the Taizishan forest farm of Hubei Province. The Pinus massoniana (Masson pine) is a common species distributed in Central China.Figure 1Geographic location of the study area. Maps were generated using ArcGIS 10.8 for Desktop (http://www.esri.com/software/arcgis/arcgis-for-desktop).Full size imageExperiment designWe chose the Pinus massoniana forest with 47a in the study area as the research object. In the typical Pinus massoniana forest, the separate layers of litter (semi-decomposed and non-decomposed layers) were collected from several 1 m × 1 m quadrat and placed in grid bags. The litter of the semi-decomposed layer have no complete outline, and the color was brown. As the litter leaves of the completely decomposed layer are powdery and are combined with the soil layer, this layer is difficult to collect. Before testing, it was necessary to clean the soil off the pine needles and then allow the litter to dry naturally. The characteristics of the semi-decomposed and non-decomposed litter layers are shown in Table 1. The soil samples need to be dried and screened by 10 mm. When filling the soil trough, every 0.1 m of soil thickness was one layer, for a total of four layers (0.4 m). The characteristics by soil particle sizes are different (Fig. 2). The soil samples were dried naturally, crushed, and then sieved. The soil trough (2 m long, 0.5 m wide and 0.5 m deep) was filled to have a bulk density of 1.53 g·m−3. In this process, an appropriate amount of water was sprinkled on the surface of each soil layer to achieve a soil moisture content consistent with the surrounding, undisturbed, or natural, state. The simulation experiment was conducted in the Jiufeng rainfall laboratory at Beijing Forestry University, China. We used a rainfall simulation system (QYJY-503T, Qingyuan Measurement Technology, Xi’an, China) used a rotary downward spray nozzle. The system is able to simulate a wide range of rainfall intensities (10 to 300 mm h−1) using various water pressure and nozzle sizes controlled by a computer system.Table 1 Characteristics of the non-decomposed and semi-decomposed layers of Pinus massoniana litter.Full size tableFigure 2Soil particle composition of study area soil layers.Full size imageAccording to the results of the field forest investigation, the litter was covered with the experimental treatments shown in Table 2. The treatments mass coverage of non-decomposed litter layer was named as follows: N1 denoted litter mass coverage 0 g·m−2, N2 was ‘the non-decomposed litter mass coverage 100 g·m−2’, N3 was ‘the non-decomposed litter mass coverage 200 g·m−2’, and N4 was ‘the non-decomposed litter mass coverage 400 g·m−2’, N5 was ‘the semi-decomposed litter mass coverage 100 g·m−2’, N6 was ‘the non-decomposed litter mass coverage 100 g·m−2 and the semi-decomposed litter mass coverage 100 g·m−2’, N7 was ‘the non-decomposed litter mass coverage 200 g·m−2 and the semi-decomposed litter mass coverage 100 g·m−2’. N2, N3 and N4 were the undissolved state of litter layer, and N4 (non-decomposed state, ND), N7 (initial stage of litter decomposition, ID), N6 (middle stage of litter decomposition, MD) and N5 (final stage of litter decomposition, FD) respectively represent different stages of litter decomposition.Table 2 The experimental design of this study.Full size tableAccording to the rainfall in the Taizishan area of Hubei Province, erosive rainfall and extreme rainstorms were selected as the research conditions. Summer rainfall events occur mainly in the summer in this area, and a rainfall intensity of 60 mm·h−1 was the most common erosive rainfall intensity. Under extreme weather conditions, the rainfall intensity can reach up to 120 mm·h−1. Our experiments were conducted with 60 and 120 mm·h−1 rain intensities with a rainfall that lasted 1 h. According to the field investigation data of forest land, this area is a low mountain and hilly area with a slope mostly between 5° and 10°. Therefore, 5° and 10° were selected for the slope treatments in this study. The combination of slope and rainfall intensity was named as follows: T1 denoted ‘Slope 5° and rainfall intensity 60 mm·h−1’, T2 was ‘Slope 10° and rainfall intensity 60 mm·h−1’, T3 was ‘Slope 5° and rainfall intensity 120 mm·h−1’, and T4 was ‘Slope 10° and rainfall intensity 120 mm·h−1’. With two rainfall intensities, two slopes, seven litter coverage gradient and two repetitions combined, this study had a total of 56 rainfall events.Experimental procedureBefore the test, the soil samples were wetted for 10 h and then drained for 2 h to eliminate the effect of the initial soil moisture on the soil detachment measurement. When the simulated rainfall started, all the runoff and sediment produced from plot were collected every 5 min in the first 10 min, and then collected once every 10 min during the subsequent 50 min. At the same time, runoff velocity, depth and temperature were measured and vernier calliper (accuracy 0.02 mm) respectively.The overland flow velocity was measured using dying method (KMnO4 solution)32. After judging the flow pattern, we confirmed the correction coefficient K value (in laminar flow state, K = 0.67; transition flow state, K = 0.70; turbulent flow state, K = 0.8). The average velocity of overland flow was obtained by multiplying the correction coefficient K and the instantaneous velocity. Runoff depth was measured using vernier calliper (accuracy 0.02 mm). Runoff temperature was measured using thermometer. When the rainfall experiment finished, the collected runoff samples were measured volumetric cylinder and then settled for at least 12 h. The clear water was decanted, and the samples were put into an oven to dry for 24 h under 105 °C. The sediment sample was dried and weighed with an electronic scale.Calculation of hydrodynamic parametersOverland flow has the characteristics of a thin water layer, large fluctuations of the underlying surface, and unstable flow velocity. At present, most scholars use open-channel flow theory to study overland flow33,34. In open-channel flow theory, the Reynold’s number (Re), Froude constant (Fr), flow index (m), resistance coefficient (f), and soil separation rate (({D}_{r})) are the basic parameters of overland flow dynamics, through Reynold’s number (Re), Froude constant (Fr), flow index (m) can distinguish flow patterns. Re is calculated as:$$Re=Rcdot V/nu ,$$where Re is the Reynolds number of the water flow, which is dimensionless, and can be used to judge the flow state of overland flow. When Re ≤ 500, the flow pattern is laminar; when 500   5000, the flow pattern is turbulent. R is the hydraulic radius (m), which is generally replaced by flow depth as measured by a vernier calliper (accuracy 0.02 mm). (V) is the average velocity (m·s−1); (nu) is the kinematic viscosity coefficient (m2·s−1), and the calculation formula is (nu) = 0.01775·10−4·(1 + 0.0337 t + 0.00021 t2), where t is the test overland flow temperature35.Fr is the Froude constant, which is the ratio of the inertial force to gravity and can be used to distinguish overland flow as rapid flow, slow flow, or critical flow. When Fr  1, the fluid is rapid flow.Fr is calculated as:$$Fr=V/sqrt{gcdot R},$$where (Fr) is the Froude constant of the water flow, which is dimensionless; (V) is the average velocity (m·s−1); g is the acceleration of gravity and has a constant value of 9.8 m·s−2; R is a hydraulic radius (m), and is generally replaced by flow depth as measured by a vernier calliper (accuracy 0.02 mm).Regression fitting is made for runoff depth (h) and single width flow (Q). The runoff depth equation for slope is as follows:$$h=k{q}^{m},$$where q is the single width flow (L·m−1·s−1); h is the depth of water on the slope (m); and m is the flow index, which reflects the turbulent characteristics of the flow state. The larger m is, the more energy the flow consumes in the work of resistance. The comprehensive index (k) reflects the characteristics of the underlying surface and the water viscosity of the slope flow. The larger k is, the stronger the surface material of the slope works on the flow.The resistance of overland flow reflects the inhibition effect of different underlying surface conditions on the velocity of overland flow. The Darcy–Weisbach formula is widely used in research because of its two advantages: applicability and dimensionlessness under laminar and turbulent flow conditions36,37.The resistance coefficient (f) is calculated as follows:$$f=8cdot gcdot Rcdot J/{V}^{2},$$where the resistance coefficient f has no dimension; g is the acceleration of gravity and is always 9.8 m·s−2; R is a hydraulic radius (m), generally replaced by flow depth measured by a vernier calliper (accuracy 0.02 mm); (V) is the average velocity (m·s−1); and J is the hydraulic gradient, which can be converted by the gradient in a uniform flow state and is generally replaced by the sine value of the gradient.Shear stress ((tau)) is the main driving force that affects the stripping of soil particles from the surface soil38. Shear stress is calculated as:$$tau =rcdot gcdot Rcdot J,$$where (tau) is the shear force of runoff (Pa); and r is the density of water and sediment concentration flow (kg·m−3). This study used a muddy water mass and volume ratio in the unseparated state to calculate the density of water and sediment concentration flow.Flow power (W) is the runoff power per unit area of water and refers to the power consumed by the weight of water acting on the riverbed surface to transport runoff and sediment. W is calculated as:$$W=tau cdot V,$$where W is the flow power (N·m−1·s−1); and (tau) is the shear force of runoff (Pa).Soil separation rate (({D}_{r})) refers to the quality of soil in which soil particles are separated from the soil per unit time. The calculation formula is as follows:$${D}_{r}={W}_{d}-{W}_{w}/tcdot A,$$where ({D}_{r}) is the rate of soil separation (kg·m−2·s−1); ({W}_{w}) is the dry weight of soil before the test; ({W}_{d}) is the dry weight of soil after the test, measured by the drying method (kg); t is the scouring time (s); and A is the surface area of the soil sample (m2). More

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    The impact of the first United Kingdom COVID-19 lockdown on environmental air pollution, digital display device use and ocular surface disease symptomatology amongst shielding patients

    Knight, H. et al. Impacts of the COVID-19 Pandemic and Self-Isolation on Students and Staff in Higher Education: A Qualitative Study. Int. J. Environ. Res. Public Health 18, 10675 (2021).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Higham, J. E., Ramírez, C. A., Green, M. A. & Morse, A. P. UK COVID-19 lockdown: 100 days of air pollution reduction? Air Quality. Atmosphere & Health https://doi.org/10.1007/s11869-020-00937-0 (2020).Article 

    Google Scholar 
    Office, P. M. s. Slides and datasets to accompany coronavirus press conference. (2020).Organization, W. H. WHO global air quality guidelines: particulate matter (PM2. 5 and PM10), ozone, nitrogen dioxide, sulfur dioxide and carbon monoxide: executive summary. (2021).Singh, A. et al. Impacts of emergency health protection measures upon air quality, traffic and public health: evidence from Oxford UK. Environ. Pollut. 293, 118584. https://doi.org/10.1016/j.envpol.2021.118584 (2022).Article 
    CAS 
    PubMed 

    Google Scholar 
    Shi, Z. et al. Abrupt but smaller than expected changes in surface air quality attributable to COVID-19 lockdowns. Science Advances 7, eabd6696, doi:doi:https://doi.org/10.1126/sciadv.abd6696 (2021).Lee, J. D., Drysdale, W. S., Finch, D. P., Wilde, S. E. & Palmer, P. I. UK surface NO2 levels dropped by 42% during the COVID-19 lockdown: impact on surface O3. Atmos. Chem. Phys. 20, 15743–15759. https://doi.org/10.5194/acp-20-15743-2020 (2020).Article 
    CAS 

    Google Scholar 
    Shi, Z. et al. Abrupt but smaller than expected changes in surface air quality attributable to COVID-19 lockdowns. Science Advances 7, eabd6696, doi:https://doi.org/10.1126/sciadv.abd6696 (2021).Ropkins, K. & Tate, J. E. Early observations on the impact of the COVID-19 lockdown on air quality trends across the UK. Sci. Total Environ. 754, 142374. https://doi.org/10.1016/j.scitotenv.2020.142374 (2021).Article 
    CAS 
    PubMed 

    Google Scholar 
    Nwanaji-Enwerem, J. C., Allen, J. G. & Beamer, P. I. Another invisible enemy indoors: COVID-19, human health, the home, and United States indoor air policy. J Expo Sci Environ Epidemiol 30, 773–775. https://doi.org/10.1038/s41370-020-0247-x (2020).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Rasha, A., Karan Jetly, J. & Shqran, S. Indoor Air Quality Monitoring Systems: A Comprehensive Review of Different IAQM Systems. International Journal of Knowledge-Based Organizations (IJKBO) 11, 1–14, doi:https://doi.org/10.4018/ijkbo.2021070101 (2021).World Health Organization. Regional Office for, E. WHO guidelines for indoor air quality: selected pollutants. xxv, 454 p. (World Health Organization. Regional Office for Europe, 2010).Stafoggia, M. et al. Long-term exposure to ambient air pollution and incidence of cerebrovascular events: Results from 11 European cohorts within the ESCAPE project. Environ. Health Perspect 122, 919–925. https://doi.org/10.1289/ehp.1307301 (2014).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Brook, R. D. et al. Particulate matter air pollution and cardiovascular disease: An update to the scientific statement from the American heart association. Circulation 121, 2331–2378. https://doi.org/10.1161/CIR.0b013e3181dbece1 (2010).Article 
    CAS 
    PubMed 

    Google Scholar 
    Raaschou-Nielsen, O. et al. Air pollution and lung cancer incidence in 17 European cohorts: prospective analyses from the European study of cohorts for air pollution effects (ESCAPE). Lancet Oncol. 14, 813–822. https://doi.org/10.1016/s1470-2045(13)70279-1 (2013).Article 
    PubMed 

    Google Scholar 
    Guan, W. J., Zheng, X. Y., Chung, K. F. & Zhong, N. S. Impact of air pollution on the burden of chronic respiratory diseases in China: Time for urgent action. Lancet 388, 1939–1951. https://doi.org/10.1016/s0140-6736(16)31597-5 (2016).Article 
    PubMed 

    Google Scholar 
    Atkinson, R. W. et al. Acute effects of particulate air pollution on respiratory admissions: Results from APHEA 2 project. Air pollution and health: A European approach. Am. J. Respir. Crit. Care Med. 164, 1860–1866. https://doi.org/10.1164/ajrccm.164.10.2010138 (2001).Article 
    CAS 
    PubMed 

    Google Scholar 
    Stapleton, F. et al. TFOS DEWS II epidemiology report. Ocular Surf. 15, 334–365. https://doi.org/10.1016/j.jtos.2017.05.003 (2017).Article 

    Google Scholar 
    Starr, C. E. et al. Dry eye disease flares: A rapid evidence assessment. Ocul. Surf. 22, 51–59. https://doi.org/10.1016/j.jtos.2021.07.001 (2021).Article 
    PubMed 

    Google Scholar 
    Torricelli, A. A. et al. Correlation between signs and symptoms of ocular surface dysfunction and tear osmolarity with ambient levels of air pollution in a large metropolitan area. Cornea 32, e11-15. https://doi.org/10.1097/ICO.0b013e31825e845d (2013).Article 
    PubMed 

    Google Scholar 
    Hwang, S. H. et al. Potential importance of ozone in the association between outdoor air pollution and dry eye disease in South Korea. JAMA Ophthalmol. 134, 503–510. https://doi.org/10.1001/jamaophthalmol.2016.0139 (2016).Article 
    PubMed 

    Google Scholar 
    Wiwatanadate, P. Acute air pollution-related symptoms among residents in Chiang Mai Thailand. J. Environ. Health 76, 76–84 (2014).CAS 
    PubMed 

    Google Scholar 
    Alves, M., Novaes, P., Morraye Mde, A., Reinach, P. S. & Rocha, E. M. Is dry eye an environmental disease? Arq. Bras. Oftalmol. 77, 193–200 https://doi.org/10.5935/0004-2749.20140050 (2014).Bourcier, T. et al. Effects of air pollution and climatic conditions on the frequency of ophthalmological emergency examinations. Br. J. Ophthalmol. 87, 809–811. https://doi.org/10.1136/bjo.87.7.809 (2003).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Hao, R. et al. Impact of air pollution on the ocular surface and tear cytokine levels: A multicenter prospective cohort study. Front. Med. (Lausanne) 9, 909330. https://doi.org/10.3389/fmed.2022.909330 (2022).Article 
    PubMed 

    Google Scholar 
    Vehof, J., Snieder, H., Jansonius, N. & Hammond, C. J. Prevalence and risk factors of dry eye in 79,866 participants of the population-based lifelines cohort study in the Netherlands. Ocul. Surf. 19, 83–93. https://doi.org/10.1016/j.jtos.2020.04.005 (2021).Article 
    PubMed 

    Google Scholar 
    Wolffsohn, J. S. et al. Demographic and lifestyle risk factors of dry eye disease subtypes: A cross-sectional study. Ocul. Surf. 21, 58–63. https://doi.org/10.1016/j.jtos.2021.05.001 (2021).Article 
    PubMed 

    Google Scholar 
    Núñez-Álvarez, C. & Osborne, N. N. Enhancement of corneal epithelium cell survival, proliferation and migration by red light: Relevance to corneal wound healing. Exp. Eye Res. 180, 231–241. https://doi.org/10.1016/j.exer.2019.01.003 (2019).Article 
    CAS 
    PubMed 

    Google Scholar 
    Marek, V. et al. Blue light phototoxicity toward human corneal and conjunctival epithelial cells in basal and hyperosmolar conditions. Free Radic. Biol. Med. 126, 27–40. https://doi.org/10.1016/j.freeradbiomed.2018.07.012 (2018).Article 
    CAS 
    PubMed 

    Google Scholar 
    Talens-Estarelles, C., García-Marqués, J. V., Cerviño, A. & García-Lázaro, S. Determining the best management strategy for preventing short-term effects of digital display use on dry eyes. Eye Contact Lens 48, 416–423. https://doi.org/10.1097/icl.0000000000000921 (2022).Article 
    PubMed 

    Google Scholar 
    GOV.UK. COVID-19: guidance on protecting people defined on medical grounds as extremely vulnerable, (2020).Joy, M. et al. Reorganisation of primary care for older adults during COVID-19: A cross-sectional database study in the UK. Br. J. Gen. Pract. 70, e540–e547. https://doi.org/10.3399/bjgp20X710933 (2020).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Schiffman, R. M., Christianson, M. D., Jacobsen, G., Hirsch, J. D. & Reis, B. L. Reliability and validity of the ocular surface disease index. Arch. Ophthalmol. 118, 615–621. https://doi.org/10.1001/archopht.118.5.615 (2000).Article 
    CAS 
    PubMed 

    Google Scholar 
    Amparo, F. & Dana, R. Web-based longitudinal remote assessment of dry eye symptoms. Ocul. Surf. 16, 249–253. https://doi.org/10.1016/j.jtos.2018.01.002 (2018).Article 
    PubMed 

    Google Scholar 
    Inomata, T. et al. Characteristics and risk factors associated with diagnosed and undiagnosed symptomatic dry eye using a smartphone application. JAMA Ophthalmol. 138, 58–68. https://doi.org/10.1001/jamaophthalmol.2019.4815 (2020).Article 
    PubMed 

    Google Scholar 
    Toth, M. & Jokić-Begić, N. Psychological contribution to understanding the nature of dry eye disease: A cross-sectional study of anxiety sensitivity and dry eyes. Health Psychol. Behav. Med. 8, 202–219. https://doi.org/10.1080/21642850.2020.1770093 (2020).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Mehra, D. & Galor, A. Digital screen use and dry eye: A review. Asia-Pacific J. Ophthalmol. 9, 491–497. https://doi.org/10.1097/apo.0000000000000328 (2020).Article 

    Google Scholar 
    Galor, A., Kumar, N., Feuer, W. & Lee, D. J. Environmental factors affect the risk of dry eye syndrome in a United States veteran population. Ophthalmology 121, 972–973. https://doi.org/10.1016/j.ophtha.2013.11.036 (2014).Article 
    PubMed 

    Google Scholar 
    Courtin, R. et al. Prevalence of dry eye disease in visual display terminal workers: A systematic review and meta-analysis. BMJ Open 6, e009675. https://doi.org/10.1136/bmjopen-2015-009675 (2016).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Torricelli, A. A. et al. Effects of ambient levels of traffic-derived air pollution on the ocular surface: Analysis of symptoms, conjunctival goblet cell count and mucin 5AC gene expression. Environ. Res. 131, 59–63. https://doi.org/10.1016/j.envres.2014.02.014 (2014).Article 
    CAS 
    PubMed 

    Google Scholar 
    Gupta, S. K., Gupta, V., Joshi, S. & Tandon, R. Subclinically dry eyes in urban Delhi: An impact of air pollution?. Ophthalmologica 216, 368–371. https://doi.org/10.1159/000066183 (2002).Article 
    CAS 
    PubMed 

    Google Scholar 
    Berg, E. J. et al. Climatic and environmental correlates of dry eye disease severity: A report from the dry eye assessment and management (DREAM) study. Trans. Vision Sci. Technol. 9, 25–25. https://doi.org/10.1167/tvst.9.5.25 (2020).Article 

    Google Scholar 
    Lang, S.-J., Abel, G. A., Mant, J. & Mullis, R. Impact of socioeconomic deprivation on screening for cardiovascular disease risk in a primary prevention population: A cross-sectional study. BMJ Open 6, e009984. https://doi.org/10.1136/bmjopen-2015-009984 (2016).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Denniston, A. K. et al. United Kingdom diabetic retinopathy electronic medical record (UK DR EMR) users group: Report 4, real-world data on the impact of deprivation on the presentation of diabetic eye disease at hospital services. Br. J. Ophthalmol. 103, 837–843. https://doi.org/10.1136/bjophthalmol-2018-312568 (2019).Article 
    PubMed 

    Google Scholar 
    Nessim, M., Denniston, A. K., Nolan, W., Holder, R. & Shah, P. Research into Glaucoma and Ethnicity (ReGAE) 8: Is there a relationship between social deprivation and acute primary angle closure?. Br. J. Ophthalmol. 94, 1304–1306. https://doi.org/10.1136/bjo.2009.160721 (2010).Article 
    PubMed 

    Google Scholar 
    Sharma, H. E. et al. The role of social deprivation in severe neovascular age-related macular degeneration. Br. J. Ophthalmol. 98, 1625–1628. https://doi.org/10.1136/bjophthalmol-2014-304959 (2014).Article 
    PubMed 

    Google Scholar 
    Bo, M., Salizzoni, P., Clerico, M. & Buccolieri, R. Assessment of indoor-outdoor particulate matter air pollution: A review. Atmosphere 8, 136 (2017).Article 

    Google Scholar 
    Strøm-Tejsen, P., Zukowska, D., Fang, L., Space, D. R. & Wyon, D. P. Advantages for passengers and cabin crew of operating a gas-phase adsorption air purifier in 11-h simulated flights. Indoor Air 18, 172–181. https://doi.org/10.1111/j.1600-0668.2007.00511.x (2008).Article 
    CAS 
    PubMed 

    Google Scholar 
    Mandell, J. T., Idarraga, M., Kumar, N. & Galor, A. Impact of air pollution and weather on dry eye. J. Clin. Med. https://doi.org/10.3390/jcm9113740 (2020).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Navarro, D. Learning Statistics with R. (Daniel Joseph Navarro, 2015). More

  • in

    A global roadmap to seize the opportunities of healthy longevity

    Building from this background the NAM took on these issues as its first-ever grand challenge, as a critical issue of import and urgency for us all. In 2018, the NAM empaneled an international, independent and multidisciplinary commission to create a global roadmap for healthy longevity, complete with evidence-based, targeted and actionable recommendations to move societies forward from an almost-exclusive focus on ‘coping with aging populations’ toward enabling individuals and societies to age successfully, and to reap the economic and societal benefits of longevity. The commission offers a way forward for governments and societies by beginning with recommendations for the next five years, and how these solutions can be financially sustainable through the creation of a virtuous cycle.To support these goals, the commission was to “(1) comprehensively address the challenges and opportunities presented by global aging population; (2) catalyze breakthrough ideas and research that will extend the human healthspan; and (3) generate transformative and scalable innovations world wide”8. The resulting comprehensive report, which was delayed in good measure by the COVID-19 pandemic, was released in June 2022 (ref. 8). We report here a summary of the high-level vision, goals, findings and recommendations of this global roadmap.The evidence for opportunities of longevity and the costs of inactionWe are seeing longer lives with increasing years spent in ill health (that is, the decompression of morbidity)9. The implications of longevity without health are costly ones for the individual, their families and for society. By contrast, scientific evidence shows that the majority of chronic diseases are preventable, and that prevention works at every age and stage of life. Further, the subset of individuals who are the beneficiaries of cumulative health-promoting conditions across the life course are demonstrating healthy longevity, defined as “the state in which years in good health approach the biological lifespan, with physical, cognitive and social functioning, enabling well-being across populations”8. However, only a minority of people in any country have the benefit of the necessary investments that promote health, and disparities in access to these investments across the life course are a major cause of unhealthy longevity. The costs of inaction in the face of widening disparities include the high risk of young people aging with more ill health, and the attendant costs to them and society.Further, the commission reports that when people have health and function in older age, the considerable cognitive and socioemotional capabilities and expertise that accrue with aging, and the prosocial goals of older age, constitute human and social capital assets that are unprecedented in both nature and scale. Contrary to disproven myths, workforce participation not only brings these valuable capabilities (such that intergenerational teams in the workplace are more productive and innovative than single-age-group teams), but older people working is also associated with more jobs for younger individuals10. In the USA and EU, it has been shown that older adults contribute 7% of gross domestic product (GDP) through paid work and the economic value of volunteering and caregiving11, even before opportunities are specifically expanded for the increasing older population. Societies that recognize this potential and invest to create both healthy longevity and the societal organizations and policies through which older adults can contribute to societal good will develop the opportunity for all ages to thrive. The return on investment will be to create older ages with health, function, dignity, meaning, purpose and opportunities — for those who desire it — to work longer, care for others or contribute in ways that they value to their community and future generations.The definition, principles and vision of ‘Vision 2050’ for healthy longevityThe global roadmap builds on the WHO ‘Decade of Healthy Ageing’, the UN Sustainable Development Goals for 2030 and other reports. It sets out principles for achieving healthy longevity using data and meaningful metrics to track achievement of outcomes and guide decision making. The report offers a vision empowered by the evidence: that, by 2050, societies will value the capabilities and assets of older people; all people will have the opportunity to live long lives with health and function; barriers to full participation by older people in society will have been solved; and that older people, with such health, will have the opportunity to engage in meaningful and productive activities. In turn, this societal engagement will create unprecedented social, human and economic capital, contributing to intergenerational well-being and cohesion, and to GDP.Implementing Vision 2050Accomplishing this vision demands ‘all-of-society’ intent — with aligned goals for healthy longevity and transformative action across public, private and academic sectors, and all of civil society and communities — and the implementation of evidence across the full and extending life course. Transforming only one component or sector (for example, health systems) will not be sufficient to create healthy longevity or its full opportunities. Rather, given that nations are complex systems, this vision for our future requires governmental leadership and transformation of all sectors of our complex societal system (Fig. 1).Fig. 1: Relevant actors for an all-of-society approach to healthy longevity.Healthy longevity requires government leadership and cooperation across all sectors. Adapted with permission from figure S-2 of ref. 8.Full size imageInvestment for healthy longevity — across the enabling sectors of health systems, social infrastructure and protections, the physical environment, and work and volunteering contributions — will require intentional planning and leadership to transform those components in tandem, and to resolve disrupters such as ageism, the social determinants of health and inequity, and pollution. These investments across all sectors will create the conditions for achieving healthy longevity and build new capital (human, social and economic) that will benefit all of society. As a result of these investments, society will see younger people thrive and move into a position to age with healthy longevity; those individuals who are already older will be recognized as valuable contributors to society in a ‘pay-it-forward’ stage of life. The underpinning social compact between citizens and government will support valuing each age group’s capabilities and goals, and the building of a society of well-being and cohesion across generations. This is at the center of the virtuous cycle for healthy longevity (Fig. 2)Fig. 2: The virtuous cycle of healthy longevity.Healthy longevity (top) is an outcome of a virtuous cycle, itself contributing to capital development (bottom left). Bottom right, capital (human, financial and social) supports enablers (work, physical environment, health systems and social infrastructure). The enablers propel the cycle, contributing to healthy longevity. Intentional investment for healthy longevity across all enabling sectors will create new capital that will benefit all of society. Adapted with permission from figure 1-4 of ref. 8.Full size imageGoals for initiating the transformation to healthy longevityThe commission identified the following changes that should occur from now to 2027 to start transformation of all of society, towards Vision 2050 and the creation of healthy longevity for all:

    Creating social cohesion, social engagement and addressing the social determinants of health through social infrastructure are among the most effective determinants of slowed aging and the prevention of chronic conditions across the life course. Financial security in older age is essential for all.

    Governments, the private sector and civil society should partner to design physical environments and infrastructure that are user-centered, and function as cohesion-enabling intergenerational communities for healthy longevity. Initiatives should focus on the inclusion of older people in the design, creating public spaces that promote social cohesion and intergenerational connection as well as mobility, physical activity and access to food, transportation, social services and engagement.

    By 2027, governments should develop strategies and plans to arrive at adequately sized, geriatrically knowledgeable public health, clinical and long-term care workforces, and an integration of the pillars of the health system and social services. Together, these dimensions would foster and extend years of good health and support the diverse health needs and well-being of older people.

    Governments should work to build the dividend of health longevity in collaboration with the business sector and civil society, to develop policies, incentives, and supportive systems that enable and encourage lifelong learning, and greater opportunities and necessary skills to engage in meaningful work or community volunteering across the lifespan.

    We summarize the commission’s recommended goals for each of these sectors in brief in Box 1. Across all sectors, the key first steps that the commission identified are ones that can resolve obstacles to change and plan the change needed to shift multiple complex systems through both top-down and bottom-up approaches, in ways appropriate to each country and context. These initiatives should create enough momentum to foster early returns on investment and optimism to propel sustained investment for subsequent stages. This would need to begin for all governments by 2023, establishing calls to action to develop and implement data-driven, all-of-society plans to build the systems, policies, organizations and infrastructure needed, and for tracking change.Box 1 Goals for 2022–2027 to initiate the transformation to healthy longevityThese goals are reproduced from Global Roadmap for Healthy Longevity8.
    Social infrastructure

    Develop evidence-based multipronged strategies to reduce ageism against all groups.

    Develop plans for ensuring basic financial security for all older people.

    Develop strategies to increase financial literacy and mechanisms for promoting working longer, pension options and savings over the life course.

    Plan opportunities for purposeful and meaningful engagement by older people at the family, community and societal levels.

    Physical environment

    At the societal level, improve broadband accessibility to reduce the digital divide and develop public transportation solutions that address first- and last-mile transportation.

    At the city level, implement mitigation strategies to reduce the negative effects of the physical environment and related emergencies on older people (for example, air pollution and climate-induced events, including extreme heat and flooding) and design environments for connection and cohesion.

    At the neighborhood level, promote and measure innovative policy solutions for healthy longevity, including affordable housing and intergenerational living, zoning and design for connection and cohesion, and the enabling of social capital.

    At the home level, update physical infrastructure and policies to address affordability, provide coliving arrangements that match people’s goals and needs, and resolve insufficiencies and inefficiencies in housing stock.

    Health systems

    Establish healthy longevity as a major goal.

    Increase investments in public health systems, which are needed to promote health and prevent disease, disability and injury at the population level, across the full life course. This may require rebalancing investments between this type of public health and medical care, recognizing that such public health is a public good and, as such, tends to be underinvested in.

    Provide adequate primary care that includes preventive screening, addresses risk factors for chronic conditions and promotes positive health behaviors, and offers a continuum of medical care, including geriatrically knowledgeable care for older adults.

    Make culturally sensitive, person-centered and equitable long-term care systems available, which (to the degree possible) offer dignity and honor people’s preferences about care settings.

    Building the healthy longevity dividend

    Governments, in collaboration with the business sector and civil society, should design (1) work environments and develop new policies that enable and encourage older adults who want or need to remain in the work force longer, and (2) engagement opportunities that strengthen communities at every stage of life.

    Governments, employers and educational institutions should prioritize redesigning education systems to support lifelong learning and training, and invest in the science of learning and training for middle-aged and older adults.

    Pilot innovations that incentivize and allow middle-aged and older adults to retool for multiple careers and/or participate as volunteers across their lifespan in roles with meaning and purpose. More

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    Multi-proxy dentition analyses reveal niche partitioning between sympatric herbivorous dinosaurs

    Versluys, J. Die Kaubewegungen von Trachodon. Palaontol. Z. 4, 80–87 (1922).
    Google Scholar 
    Kripp, D. Die Kaubewegung und Lebensweise von Edmontosaurus spec. auf Grund der mechanischkonstruktiven analyse. Palaeobiologica 5, 409–422 (1933).
    Google Scholar 
    Ostrom, J. H. Cranial morphology of the hadrosaurian dinosaurs of North America. Bull. Am. Mus. Nat. Hist. 122, 39–186 (1961).
    Google Scholar 
    Ostrom, J. H. A functional analysis of jaw mechanics in the dinosaur. Triceratops. Postilla. 88, 1–35 (1964).MathSciNet 

    Google Scholar 
    Galton, P. M. The cheeks of ornithischian dinosaurs. Lethaia 6, 67–89. https://doi.org/10.1111/j.1502-3931.1973.tb00873.x (1973).Article 

    Google Scholar 
    Galton, P. M. Herbivorous adaptations of Late Triassic and Early Jurassic dinosaurs. In The Beginning of the Age of Dinosaurs (ed. Padian, K.) 203–221 (Cambridge University Press, 1986).
    Google Scholar 
    Weishampel, D. B. Hadrosaurid jaw mechanics. Acta Palaeontol. Pol. 28, 271–280 (1983).
    Google Scholar 
    Weishampel, D. B. The evolution of jaw mechanisms in ornithopod dinosaurs. Adv. Anat. Embryol. Cell. Biol. 87, 1–2 (1984).Article 
    CAS 
    PubMed 

    Google Scholar 
    Weishampel, D. B. Interactions between Mesozoic plants and vertebrates: fructifications and seed predation. Neues Jahrb. Geol. Paläontol. Abh. 167, 224–250 (1984).
    Google Scholar 
    Weishampel, D. B. & Norman, D. B. Vertebrate herbivory in the Mesozoic: Jaws, plants, and evolutionary metrics. In Paleobiology of the Dinosaurs Special Papers 238 (ed. Farlow, J. O.) 87–100 (Geological Society of America, 1989).Chapter 

    Google Scholar 
    Norman, D. B. & Weishampel, D. B. Feeding mechanisms in some small herbivorous dinosaurs: processes and patterns. In Biomechanics and Evolution (eds Rayner, J. M. V. & Wooton, R. J.) 161–181 (Cambridge University Press, 1991).
    Google Scholar 
    Sereno, P., Zijin, Z. & Lin, T. A new psittacosaur from Inner Mongolia and the parrot-like structure and function of the psittacosaur skull. Proc. Roy. Soc. B. 277, 199–209. https://doi.org/10.1098/rspb.2009.0691 (2010).Article 

    Google Scholar 
    Barrett, P. M. Paleobiology of herbivorous dinosaurs. Annu. Rev. Earth Planet. Sci. 42(1), 207–230. https://doi.org/10.1146/annurev-earth-042711-105515 (2014).Article 
    CAS 

    Google Scholar 
    Erickson, G. M. et al. Wear biomechanics in the slicing dentition of the giant horned dinosaur Triceratops. Sci. Adv. 1(5), e1500055. https://doi.org/10.1126/sciadv.1500055 (2015).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Nabavizadeh, A. Hadrosauroid jaw mechanics and the functionalsignificance of the predentary bone. In The hadrosaurs: Proceedings of the International Hadrosaur Symposium (eds Evans, D. & Eberth, D.) 467–482 (Indiana University Press, 2014).
    Google Scholar 
    Nabavizadeh, A. Evolutionary trends in the jaw adductor mechanics of ornithischian dinosaurs. Anat. Rec. 299(3), 271–294. https://doi.org/10.1002/ar.23306 (2016).Article 

    Google Scholar 
    Nabavizadeh, A. new reconstruction of cranial musculature in ceratopsian dinosaurs: Implications for jaw mechanics and ‘cheek’anatomy. FASEB J. 30, lb27–lb27. https://doi.org/10.1096/fasebj.30.1_supplement.lb27 (2016).Article 

    Google Scholar 
    Nabavizadeh, A. new reconstruction of cranial musculature in ornithischian dinosaurs: Implications for feeding mechanismsand buccal anatomy. Anat. Rec. 303, 347–362. https://doi.org/10.1002/ar.23988 (2020).Article 

    Google Scholar 
    Varriale, F. J. Dental microwear reveals mammal-like chewing in the neoceratopsian dinosaur Leptoceratops gracilis. PeerJ 4, e2132. https://doi.org/10.7717/peerj.2132 (2016).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Melstrom, K. M., Chiappe, L. M. & Smith, N. D. Exceptionally simple, rapidly replaced teeth in sauropod dinosaurs demonstrate a novel evolutionary strategy for herbivory in Late Jurassic ecosystems. BMC Evol. Biol. 21(1), 1–12. https://doi.org/10.1186/s12862-021-01932-4 (2021).Article 

    Google Scholar 
    Norman, D. B. On the cranial morphology and evolution of ornithopod dinosaurs. Proc. Zool. Soc. Lond. 52, 521–547 (1984).
    Google Scholar 
    Norman, D. B. & Weishampel, D. B. Ornithopod feeding mechanisms: Their bearing on the evolution of herbivory. Am. Nat. 126, 151–164. https://doi.org/10.1086/284406 (1985).Article 

    Google Scholar 
    Norman, D. B. & Weishampel, D. B. Vegetarian dinosaurs chew it differently-living mammals can chew plants for more effectively than reptiles. Yet some dinosaurs were surprisingly adept chewers. This unexpected ability may have been crucial in their evolution. New Sci. 114(1559), 42–45 (1987).
    Google Scholar 
    Rybczynski, N., Tirabasso, A., Bloskie, P., Cuthbertson, R. & Holliday, C. A three-dimensional animation model of Edmontosaurus (Hadrosauridae) for testing chewing hypotheses. Palaeontol. Electron. 11(2), 9A (2008).
    Google Scholar 
    Williams, V. S., Barrett, P. M. & Purnell, M. A. Quantitative analysis of dental microwear in hadrosaurid dinosaurs, and the implications for hypotheses of jaw mechanics and feeding. PNAS 106(27), 11194–11199. https://doi.org/10.1073/pnas.0812631106 (2009).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Cuthbertson, R. S., Tirabasso, A., Rybczynski, N. & Holmes, R. B. Kinetic limitations of intracranial joints in Brachylophosaurus canadensis and Edmontosaurus regalis (Dinosauria: Hadrosauridae), and their implications for the chewing mechanics of hadrosaurids. Anat. Rec. 295, 968–979. https://doi.org/10.1002/ar.22458 (2012).Article 

    Google Scholar 
    Erickson, G. M. & Zelenitsky, D. K. Osteohistology and occlusal morphology of Hypacrosaurus stebengeri teeth throughout ontogeny with comments on wear-induced form and function. In Hadrosaurs (eds Eberth, D. A. & Evans, D. C.) 422–432 (Indiana University Press, 2014).
    Google Scholar 
    Barrett, P. M. Tooth wear and possible jaw action of Scelidosaurus harrisonii Owen and a review of feeding mechanisms in other thyreophoran dinosaurs. In The Armored Dinosaurs (ed. Carpenter, K.) 25–52 (Indiana University Press, 2001).
    Google Scholar 
    Rybczynski, N. & Vickaryous, M. K. Evidence of complex jaw movement in the Late Cretaceous ankylosaurid Euoplocephalus tutus (Dinosauria: Thyreophora). In The Armored Dinosaurs (ed. Carpenter, K.) 299–317 (Indiana University Press, 2001).
    Google Scholar 
    Mallon, J. C. & Anderson, J. S. The functional and palaeoecological implications of tooth morphology and wear for the megaherbivorous dinosaurs from the Dinosaur Park Formation (Upper Campanian) of Alberta, Canada. PLoS ONE 9(6), e98605. https://doi.org/10.1371/journal.pone.0098605 (2014).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Mallon, J. C. & Anderson, J. S. Implications of beak morphology for the evolutionary paleoecology of the megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada. Palaeogeogr. Palaeoclimatol. Palaeoecol. 394, 29–41. https://doi.org/10.1016/j.palaeo.2013.11.014 (2014).Article 

    Google Scholar 
    Ősi, A., Barrett, P. M., Földes, T. & Tokai, R. Wear pattern, dental function, and jaw mechanism in the Late Cretaceous ankylosaur Hungarosaurus. Anat. Rec. 297(7), 1165–1180. https://doi.org/10.1002/ar.22910 (2014).Article 

    Google Scholar 
    Ősi, A., Prondvai, E., Mallon, J. & Bodor, E. R. Diversity and convergences in the evolution of feeding adaptations in ankylosaurs (Dinosauria: Ornithischia). Hist. Biol. 29(4), 539–570. https://doi.org/10.1080/08912963.2016.1208194 (2017).Article 

    Google Scholar 
    Hill, R. V., D’Emic, M. D., Bever, G. S. & Norell, M. A. A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia. Zool. J. Linn. Soc. 175(4), 892–909. https://doi.org/10.1111/zoj.12293 (2015).Article 

    Google Scholar 
    Lautenschlager, S., Brassey, C. A., Button, D. J. & Barrett, P. M. Decoupled form and function in disparate herbivorous dinosaur clades. Sci. Rep. 6(1), 1–10. https://doi.org/10.1038/srep26495 (2016).Article 
    CAS 

    Google Scholar 
    Skutschas, P. P. et al. Wear patterns and dental functioning in an Early Cretaceous stegosaur from Yakutia, Eastern Russia. PLoS ONE 16(3), e0248163. https://doi.org/10.1371/journal.pone.0248163 (2021).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Strickson, E., Prieto-Márquez, A., Benton, M. J. & Stubbs, T. L. Dynamics of dental evolution in ornithopod dinosaurs. Sci. Rep. 6, 28904. https://doi.org/10.1038/srep28904 (2016).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Virág, A. & Ősi, A. Morphometry, microstructure, and wear pattern of neornithischian dinosaur teeth from the Upper Cretaceous Iharkút locality (Hungary). Anat. Rec. 300(8), 1439–1463. https://doi.org/10.1002/ar.23592 (2017).Article 

    Google Scholar 
    Mallon, J. C. & Anderson, J. S. Skull ecomorphology of megaherbivorous dinosaurs from the Dinosaur Park Formation (Upper Campanian) of Alberta, Canada. PLoS ONE 8(7), e67182. https://doi.org/10.1371/journal.pone.0067182 (2013).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Botfalvai, G., Ősi, A. & Mindszenty, A. Taphonomic and paleoecologic investigations of the Late Cretaceous (Santonian) Iharkút vertebrate assemblage (Bakony Mts, northwestern Hungary). Palaeogeogr. Palaeoclimatol. Palaeoecol. 417, 379–405. https://doi.org/10.1016/j.palaeo.2014.09.032 (2015).Article 

    Google Scholar 
    Botfalvai, G., Haas, J., Bodor, E. R., Mindszenty, A. & Ősi, A. Facies architecture and palaeoenvironmental implications of the upper Cretaceous (Santonian) Csehbánya formation at the Iharkút vertebrate locality (Bakony Mountains, Northwestern Hungary). Palaeogeogr. Palaeoclimatol. Palaeoecol. 441, 659–678. https://doi.org/10.1016/j.palaeo.2015.10.018 (2016).Article 

    Google Scholar 
    Ősi, A. et al. The Late Cretaceous continental vertebrate fauna from Iharkút, western Hungary: A review. In Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems (ed. Godefroit, P.) 532–569 (Indiana University Press, 2012).
    Google Scholar 
    Wells, N. A. Making thin sections. In Paleotechniques (eds Feldmann, R. M. et al.) 120–129 (University of Tennessee, 1989).
    Google Scholar 
    Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. NIH Image to ImageJ: 25 years of image analysis. Nat. Methods 9(7), 671–675. https://doi.org/10.1038/nmeth.2089 (2012).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Evans, A. R. Surfer Manipulator. http://evomorph.org/surfermanipulator (2011).Evans, A. R., Wilson, G. P., Fortelius, M. & Jernvall, J. High-level similarity of dentitions in carnivorans and rodents. Nature 445, 78–81. https://doi.org/10.1038/nature05433 (2007).Article 
    CAS 
    PubMed 

    Google Scholar 
    Wilson, G. P. et al. Adaptive radiation of multituberculate mammals before the extinction of dinosaurs. Nature 483, 457–460. https://doi.org/10.1038/nature10880 (2012).Article 
    CAS 
    PubMed 

    Google Scholar 
    Ungar, P. S. Dental microwear of European Miocene catarrhines: Evidence for diets and tooth use. J. Hum. Evol. 31, 355–366. https://doi.org/10.1006/jhev.1996.0065 (1996).Article 

    Google Scholar 
    Ungar, P. S. A semiautomated image analysis procedure for the quantification of dental microwear II. Scanning. 17, 57–59. https://doi.org/10.1002/sca.4950170108 (1995).Article 
    CAS 
    PubMed 

    Google Scholar 
    Ungar, P. S., Brown, C. A., Bergstrom, T. S. & Walker, A. Quantification of dental microwear by tandem scanning confocal microscopy and scale-sSensitive fractal analyses. Scanning 25, 185–193. https://doi.org/10.1002/sca.4950250405 (2003).Article 
    PubMed 

    Google Scholar 
    Ungar, P. S., Merceron, G. & Scott, R. S. Dental microwear texture analysis of Varswater bovids and Early Pliocene paleoenvironments of langebaanweg, Western Cape Province, South Africa. J. Mammal. Evol. 14, 163–181. https://doi.org/10.1007/s10914-007-9050-x (2007).Article 

    Google Scholar 
    Scott, J. R. Dental microwear texture analysis of extant African Bovidae. Mammalia 76, 157–217. https://doi.org/10.1515/mammalia-2011-0083 (2012).Article 

    Google Scholar 
    Merceron, G., Hofman-Kaminska, E. & Kowalczyk, R. 3D dental microwear texture analysis of feeding habits of sympatric ruminants in the Białowieza Primeval Forest, Poland. For. Ecol. Manag. 328, 262–269. https://doi.org/10.1016/j.foreco.2014.05.041 (2014).Article 

    Google Scholar 
    Caporale, S. S. & Ungar, P. S. Rodent incisor microwear as a proxy for ecological reconstruction. Palaeogeog. Palaeocl. Palaeoecol. 446, 225–233. https://doi.org/10.1016/j.palaeo.2016.01.013 (2016).Article 

    Google Scholar 
    R Core Team. R. A language and environment for statistical computing. R Foundation for Statistical Computing https://www.R-project.org/ (2021).Erickson, G. M. Incremental lines of von Ebner in dinosaurs and the assessment of tooth replacement rates using growth line counts. PNAS 93(25), 14623–14627. https://doi.org/10.1073/pnas.93.25.14623 (1996).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Godefroit, P. et al. Extreme tooth enlargement in a new Late Cretaceous rhabdodontid dinosaur from Southern France. Sci. Rep. 7(1), 1–9. https://doi.org/10.1038/s41598-017-13160-2 (2017).Article 
    CAS 

    Google Scholar 
    Edmund, G. Tooth replacement phenomena in the lower vertebrates. Life. Sci. Contrib. R. Ont. Mus. 52, 1–190 (1960).
    Google Scholar 
    D’Emic, M. D., Whitlock, J. A., Smith, K. M., Fisher, D. C. & Wilson, J. A. Evolution of high tooth replacement rates in sauropod dinosaurs. PLoS ONE 8(7), e69235. https://doi.org/10.1371/journal.pone.0069235 (2013).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Ősi, A., Prondvai, E., Butler, R. & Weishampel, D. B. Phylogeny, histology and inferred body size evolution in a new rhabdodontid dinosaur from the Late Cretaceous of Hungary. PLoS ONE 7(9), e44318. https://doi.org/10.1371/journal.pone.0044318 (2012).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Weishampel, D. B., Jianu, C. M., Csiki, Z. & Norman, D. B. Osteology and phylogeny of Zalmoxes (ng), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. J. Syst. Palaeontol. 1(2), 65–123. https://doi.org/10.1017/S1477201903001032 (2003).Article 

    Google Scholar 
    Melstrom, K. M. The relationship between diet and tooth complexity in living dentigerous saurians. J. Morphol. 278, 500–522 (2017).Article 
    PubMed 

    Google Scholar 
    LeBlanc, A. R. H., Reisz, R. R., Evans, D. C. & Bailleul, A. M. Ontogeny reveals function and evolution of the hadrosaurid dinosaur dental battery. BMC Evol. Biol. 16(1), 1–13. https://doi.org/10.1186/s12862-016-0721-1 (2016).Article 

    Google Scholar 
    Erickson, G. M. et al. Complex dental structure and wear biomechanics in hadrosaurid dinosaurs. Science 338(6103), 98–101. https://doi.org/10.1126/science.1224495 (2012).Article 
    CAS 
    PubMed 

    Google Scholar 
    Norman, D. B. & Weishampel, D. B. Iguanodontidae and related Ornithopoda. In The Dinosauria (eds Weishampel, D. B. et al.) 510–533 (University of California Press, 1990).
    Google Scholar 
    Hulke, J. W. An attempt at a complete osteology of Hypsilophodon foxii, a British Wealden dinosaur. Philos. Trans. R. Soc. Lond. 172, 1035–1062. https://doi.org/10.1098/rstl.1882.0025 (1882).Article 

    Google Scholar 
    Sternberg, C. H. Thescelosaurus edmontonensis, n. sp., and classification of the Hypsilophodontidae. J. Paleontol. 14, 481–494 (1940).
    Google Scholar 
    Galton, P. M. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull. Br. Mus. Nat. Hist. 25(1), 1–152 (1974).
    Google Scholar 
    Norman, D. B. On the anatomy of Iguanodon atherfieldensis (Ornithischia: Ornithopoda). Bull. Inst. Roy. Sci. Nat. Belgique 56, 281–372 (1986).
    Google Scholar 
    Norman, D. B. & Barrett, P. M. Ornithischian dinosaurs from the lower Cretaceous (Berriasian) of England. Spec. Pap. Palaeontol. 68, 161–190 (2002).
    Google Scholar 
    Kosch, J. C. & Zanno, L. E. Sampling impacts the assessment of tooth growth and replacement rates in archosaurs: Implications for paleontological studies. PeerJ 8, e9918. https://doi.org/10.7717/peerj.9918 (2020).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Janis, C. M. & Fortelius, M. On the means whereby mammals achieve increased functional durability of their dentitions with special reference to limiting factors. Biol. Rev. 63, 197–230. https://doi.org/10.1111/j.1469-185X.1988.tb00630.x (1988).Article 
    CAS 
    PubMed 

    Google Scholar 
    You, H., Ji, Q. & Li, D. Lanzhousaurus magnidens gen. et sp. nov. from Gansu Province, China: The largest-toothed herbivorous dinosaur in the world. Geol. Bull. Chi 24(9), 785–794 (2005).
    Google Scholar 
    Suarez, C. A., You, H. L., Suarez, M. B., Li, D. Q. & Trieschmann, J. B. Stable isotopes reveal rapid enamel elongation (amelogenesis) rates for the Early Cretaceous iguanodontian dinosaur Lanzhousaurus magnidens. Sci. Rep. 7, 15319. https://doi.org/10.1038/s41598-017-15653-6 (2017).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Upchurch, P. & Barrett, P. M. The evolution of sauropod feeding mechanisms. In Evolution of Herbivory in Terrestrial Vertebrates: Perspectives from the Fossil Record (ed. Sues, H. D.) 79–122 (Cambridge University Press, 2000).Chapter 

    Google Scholar 
    Sereno, P. C. & Wilson, J. A. Structure and evolution of a sauropod tooth battery in Curry. In The Sauropods: Evolution and Paleobiology (eds Rogers, K. A. & Wilson, J. A.) 157–177 (University of California Press, 2005).
    Google Scholar 
    Brown, B. & Schlaikjer, E. M. The structure and relationships of Protoceratops. Ann. N. Y. Acad. Sci. 40(3), 133–265. https://doi.org/10.1111/j.1749-6632.1940.tb57047.x (1940).Article 

    Google Scholar 
    Solounias, N., Teaford, M. & Walker, A. Interpreting the diet of extinct ruminants-the case of a non-browsing giraffid. Paleobiology 14, 287–300. https://doi.org/10.1017/S009483730001201X (1988).Article 

    Google Scholar 
    Walker, A. & Teaford, M. Inferences from quantitative analysis of dental microwear. Folia Primatol. 53, 177–189. https://doi.org/10.1159/000156415 (1989).Article 
    CAS 

    Google Scholar 
    Ungar, P. S. Mammalian dental function and wear: A review. Biosurf. Biotribol. 1(1), 25–41. https://doi.org/10.1016/j.bsbt.2014.12.001 (2015).Article 
    MathSciNet 

    Google Scholar 
    Janis, C. M. An estimation of tooth volume and hypsodonty indices in ungulate mammals, and the correlation of these factors with dietary preferences. Mém. Mus. Natl. Hist. Nat. Sér. C Géol. 53, 367–387 (1988).
    Google Scholar 
    Lucas, P. W. et al. The role of dust, grit and phytoliths in tooth wear. Ann. Zool. Fenn. 51(1–2), 143–152. https://doi.org/10.5735/086.051.0215 (2014).Article 

    Google Scholar 
    Winkler, D. E. et al. Shape, size, and quantity of ingested external abrasives influence dental microwear texture formation in guinea pigs. Proc. Nat. Acad. Sci. 117, 22264–22273. https://doi.org/10.1073/pnas.2008149117 (2020).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Kaiser, T. M. et al. Nano-indentation of native phytoliths and dental tissues: Implications for herbivore-plant combat and dental wear proxies. Evol. Syst. 2, 55–63. https://doi.org/10.3897/evolsyst.2.22678 (2018).Article 

    Google Scholar 
    Winkler, D. E. et al. Forage silica and water content control dental surface texture in guinea pigs and provide implications for dietary reconstruction. Proc. Nat. Acad. Sci. 116, 1325–1330. https://doi.org/10.1073/pnas.1814081116 (2019).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Ősi, A. & Makádi, L. New remains of Hungarosaurus tormai (Ankylosauria, Dinosauria) from the Upper Cretaceous of Hungary: Skeletal reconstruction and body mass estimation. Palaontol. Z. 83(2), 227–245. https://doi.org/10.1007/s12542-009-0017-5 (2009).Article 

    Google Scholar 
    Winkler, D. E., Schulz-Kornas, E., Kaiser, T. M. & Tütken, T. Dental microwear texture reflects dietary tendencies in extant Lepidosauria despite their limited use of oral food processing. Proc. R. Soc. B 286, 20190544. https://doi.org/10.1098/rspb.2019.0544 (2019).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Bestwick, J., Unwin, D. M., Butler, R. J. & Purnell, M. A. Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis. Nat. Commun. 11, 1–9. https://doi.org/10.1038/s41467-020-19022-2 (2020).Article 
    CAS 

    Google Scholar 
    Sakaki, H. et al. Non-occlusal dental microwear texture analysis of a titanosauriform sauropod dinosaur from the Upper Cretaceous (Turonian) Tamagawa Formation, northeastern Japan. Cret. Res. 136, 105218. https://doi.org/10.1016/j.cretres.2022.105218 (2022).Article 

    Google Scholar 
    Fiorillo, A. R. Dental microwear on the teeth of Camarasaurus and Diplodocus; implications for sauropod paleoecology. In Fifth Symposium on Mesozoic Terrestrial Ecosystems and Biota (eds Kielan-Jaworowska, Z. et al.) 23–24 (Paleontologisk Museum, 1991).
    Google Scholar 
    Mallon, J. C., Cuthbertson, R. S. & Tirabasso, A. Hadrosaurid jaw mechanics as revealed by cranial joint limitations and dental microwear analysis. In Hadrosaur Symposium Abstract Volume (eds Braman, D. R. et al.) 87–90 (Royal Tyrrell Museum of Palaeontology, 2011).
    Google Scholar 
    Fiorillo, A. R. Dental microwear patterns of the sauropod dinosaurs Camarasaurus and Diplodocus: Evidence for resource partitioning in the Late Jurassic of North America. Hist. Biol. 13, 1–16. https://doi.org/10.1080/08912969809386568 (1998).Article 

    Google Scholar 
    Sereno, P. C. et al. Structural extremes in a Cretaceous dinosaur. PLoS ONE 2(11), e1230. https://doi.org/10.1371/journal.pone.0001230 (2007).Article 
    PubMed 
    PubMed Central 

    Google Scholar 
    Whitlock, J. A. Inferences of diplodocoid (Sauropoda: Dinosauria) feeding behavior from snout shape and microwear analyses. PLoS ONE 6(4), e18304. https://doi.org/10.1371/journal.pone.0018304 (2011).Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 
    Fiorillo, A. R. Microwear patterns on the teeth of northern high latitude hadrosaurs with comments on microwear patterns in hadrosaurs as a function of latitude and seasonal ecological constraints. Palaeontol. Electron. 14(3), 20A (2011).
    Google Scholar 
    Bell, P. R., Snively, E. & Shychoski, L. A comparison of the jaw mechanics in hadrosaurid and ceratopsid dinosaurs using finite element analysis. Anat. Rec. 292(9), 1338–1351. https://doi.org/10.1002/ar.20978 (2009).Article 

    Google Scholar 
    Chin, K. & Gill, B. D. Dinosaurs, dung beetles, and conifers: Participants in a Cretaceous food web. Palaios 11, 280–285. https://doi.org/10.2307/3515235 (1996).Article 

    Google Scholar 
    Brown, C. M. et al. Dietary palaeoecology of an early Cretaceous armoured dinosaur (Ornithischia; Nodosauridae) based on floral analysis of stomach contents. Roy. Soc. Open Sci. 7(6), 200305. https://doi.org/10.1098/rsos.200305 (2020).Article 
    CAS 

    Google Scholar 
    Crane, P. C., Friis, E. M. & Pedersen, K. R. The origin and early diversification of angiosperms. Nature 374, 27–33 (1995).Article 
    CAS 

    Google Scholar 
    Friis, E. M., Crane, P. R. & Pedersen, K. R. Early Flowers and Angiosperm Evolution 1–596 (Cambridge University Press, 2011). https://doi.org/10.1017/CBO9780511980206.Book 

    Google Scholar 
    Benson, R. B., Hunt, G., Carrano, M. T. & Campione, N. Cope’s rule and the adaptive landscape of dinosaur body size evolution. Palaeontology 61, 13–48. https://doi.org/10.1111/pala.12329 (2018).Article 

    Google Scholar 
    Hummel, J. et al. In vitro digestibility of fern and gymnosperm foliage: Implications for sauropod feeing ecology and diet selection. Proc. Royal Soc. B 275, 1015–1021. https://doi.org/10.1098/rspb.2007.1728 (2008).Article 

    Google Scholar 
    Gee, C. T. Dietary options for the sauropod dinosaurs from an integrated botanical and paleobotanical perspective. In Biology of the Sauropod Dinosaurs: Understanding the Life of Giants (eds Klein, K. et al.) 34–56 (Indiana University Press, 2011).
    Google Scholar 
    Peters, R. H. The Ecological Implications of Body Size 1–329 (Cambridge University Press, 1983).Book 

    Google Scholar 
    Jarman, P. J. The social organisation of antelope in relation to their ecology. Behaviour 48, 215–267 (1974).Article 

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    Researchers who reach far beyond their disabilities

    Scientists with visible and invisible disabilities take on adversity, helping themselves and others.Shigehiro Namiki always wanted to study insects. After his PhD research at the University of Tsukuba, he was a postdoctoral fellow, then a staff scientist at Janelia Research Campus. Among his projects, Namiki worked with others on a method to analyze how the few so-called descending neurons in fruit flies control a wide range of movements and behavior. These neurons run from the brain to the ventral nerve cord and branch out to circuits that control the insect’s neck, legs and wings. More

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    Alternative stable ecological states observed after a biological invasion

    Study systemOur focal ecosystem is in Selvíria, state of Mato Grosso do Sul, Brazil ((hbox {20}^{circ }) (22′) (41.86”) S, (hbox {51}^{circ }) (24′) (58.90”) W), on a property owned by the São Paulo State University (UNESP). The location covers 350 ha of pasture composed of liverseed grass (Urochloa decumbens). The native vegetation was removed, pasture areas were implemented, and livestock was introduced in the 1970s, maintaining this configuration during the following 50 years. The climate of this area is categorized as equatorial savanna, with dry periods concentrated mostly during the winter, from April to August. During our sampling period (from November 23th, 1989, to November 19th, 2015), no vermifuges and insecticides that could affect negatively the community of dung beetles associated with cow pads were used1.The native dung beetle community at this site is composed of dwellers and tunnelers. Dwellers comprise the Aphodiinae subfamily, whereas all the tunnelers belong to the Scarabaeinae subfamily31. In total, there were eight species classified as dwellers (Ataenius crenulatus, A. picinus and Atanius aequalis-platensis grouped as one species, Blackburneus furcatus, Genieridium bidens, Labarrus pseudolividus, Nialaphodius nigrita and Trichillum externepunctatum) and ten native tunnelers (Ateuchus nr. puncticollis, A. vividus, Canthidium nr. pinotoides, Dichotomius bos, D. semiaeneus, D. sexdentatus, Ontherus appendiculatus, O. dentatus, O. sulcator). These species were chosen for our study because, as the invasive tunneler D. gazella (also from the Scarabaeinae subfamily), they all co-occur in pasture and exploit the same resource (cow pad)32. The initial establishment of D. gazella caused the loss of most of the native tunnelers from the community, with the invader becoming the overwhelming representative of the functional group, and an initial decrease of abundance for dwellers. Differently from native tunnelers, however, dwellers were able to recover their number a few years after invasion (Fig. 1a, Fig. S1).As reported in1, the abundance of dung beetles was significantly affected by both local minimum temperature and relative humidity. The influence of these two factors is expected, as they determine egg and larval survival and development of dung beetles. For example, because dung beetles are poikilotherms, environmental temperature is key to their development and fecundity33. One of the main dweller species, Labarrus pseudolividus, is widely found in locations with temperature averages ranging between (hbox {12},^{circ }hbox {C}) and (hbox {18},^{circ }hbox {C})34, making it tolerant to colder local temperatures. On the other hand, for D. gazella the lower developmental threshold is (hbox {15.5},^{circ }hbox {C}) (individuals cannot survive below this temperature), and the optimum temperature for population growth is (hbox {28},^{circ }hbox {C})35. For both groups, physiological growth and reproduction rates are maintained even when outside temperatures are close to the lower developmental threshold; dwellers, for example, live inside the dung pile, where temperature is higher and less variable than outside36,37. However, while tunnelers oviposit deep in the soil to protect the eggs, warmer and drier conditions reduce dweller egg viability on dung piles since they are exposed38. Low humidity conditions lead to drier dung and can cause egg and insect dessication. In addition, dwellers from our focal system have Palearctic evolutionary origins39; D. gazella’s natural distribution ranges from central to southern Africa40, presenting high physiological plasticity that allows it to tolerate high temperatures and low relative humidity better than other tunneler species41.Functional-group data collection and community structure characterizationDung beetles were collected once a week in a black-light flight intercept trap42, which guarantees the collection of coprophagic beetles. During all collection periods, climate variables were also collected from a meteorological station located within 2 km of our collecting site. See1 for the complete description of the collection process and database. For our purposes, we retained the species, number of individuals per species, and climate variables for each week sampled (Supplementary Information, SI, Figs. S1–S2).We focused first on the weekly abundance data, which we needed to process in order to avoid spurious results in our analyses stemming from the measurement protocol. Specifically, we filtered out seasonal low values associated with sampling in the coldest periods, when few beetles are captured because the reduced activity in all functional groups restricts their spatio-temporal distribution43. Including such samples would not be representative of the community and could bias the analysis since we are investigating community composition (i.e. proportions, very sensitive to low sampling). Thus, we considered only samples with a total number of beetles (that is, summing up all groups together) higher than the value of the median of all data, a conservative threshold that retains observations that allow for as much representation of the community as possible. As will become evident in the Results section and Supplementary Information, less conservative choices for the threshold did not alter our main conclusions.Following Mesquita -Filho et al.1, we categorized all sampled species into either dwellers or tunnelers. D. gazella is a tunneler and, as explained above, the native tunneler species experienced massive declines in abundance after its establishment, leaving D. gazella as almost the single representative in the tunneler functional group during the period of observation1. Thus, given the sharp contrast in community composition, we also separated the data into before and after invasion using to that end the 200th week, when D. gazella was first observed at the study site (September 11th, 1993, starting date for what we will call “after invasion”, our focal period henceforth).To describe community functional composition (i.e. system state) through time, we derived a normalized functional group ratio. First, because the abundance of each functional group spanned up to four orders of magnitude, we performed a logarithmic transformation of the number of captured insects from each group i, (log _{10}(N_{i}+K)), following  Yamamura44. Here, we chose (K=1), but the value of K did not alter our results qualitatively. In addition, the original data showed random mismatches in the phenology of each group, which gave the wrong impression of extreme short-term shifts in functional group dominance within the community. To avoid such artifacts, we used nonparametric local regression (LOESS)45 to smooth the dynamics of each group46. For this smoothing, we employed the loess function in the R software 3.6.147 with a smooth parameter equal to 0.25, but other moderate values (or an optimal value calculated with Bayesian inference by the R function optimal_span) did not alter our conclusions. Finally, we extracted back from the smoothed curve the number of beetles within each functional group to calculate the fraction (f_{dwell}) that measures the relative abundance of dwellers:$$begin{aligned} f_{dwell} = frac{N_D}{N_D+N_T} end{aligned}$$
    (1)
    where (N_D) corresponds to the number of dwellers per week and (N_T) corresponds to the number of native tunnelers (for the period before invasion), or only the number of D. gazella observed per week (after invasion), using their corresponding smoothed curves. Including also native tunnelers after invasion did not alter our conclusions.Climate driverWe devised a single climatic driver variable that merges the weekly measurement of temperature and relative humidity over the years, abiotic factors key to the survival and reproduction of both groups (see above). We first converted minimum temperatures and relative humidity to normalized climate variables using a min-max normalization (a feature scaling that uses the total range of temperatures or relative humidity, respectively, as normalization factor):$$begin{aligned} T = frac{T_{week} – T_{min}}{T_{max}-T_{min}};;,~ ~ ~ ~ ~ ~ RH = frac{RH_{week} – RH_{min}}{RH_{max}-RH_{min}};;, end{aligned}$$
    (2)
    where T corresponds to the normalized temperature, (T_{week}) is the weekly temperature, and (T_{max}) and (T_{min}) are the absolute maximum and minimum temperatures observed during the whole sampling period, respectively. We used a similar notation for relative humidity, RH. Based on the information above regarding beetle response to climate, the merged climate factor c was defined as the relationship:$$begin{aligned} c = frac{T}{RH};;, end{aligned}$$
    (3)
    for (RHne 0). That is, higher temperatures and/or drier conditions (expected to favor D. gazella) lead to higher values for c. On the other hand, lower temperatures and/or more humid conditions (expected to favor dwellers) imply lower values for c. Intermediate values of c can represent either moderate or extreme values for both T and RH.Identifying ecological states and quantifying resilienceWith our (f_{dwell}) data as an index of community composition (i.e. system state), we calculated kernel density functions to interpolate a continuous probability distribution of the relative fraction of dwellers in the community, (p_{n}(f_{dwell})) (function density, R software 3.1.647) for a given range of climatic driver c values. We grouped the (f_{dwell}) data using ranges for c of size 0.4, to ensure a significant amount of weekly samples that allowed for the reconstruction of these probability distributions (see Table S1, first column). Note that bins with extreme values showed few data points (see first and last rows in Table S1), and thus were rejected to prevent misleading results due to reduced sampling. Also note that, for the density function, we used the default Gaussian kernel with a smoothing bandwidth adjusted to be (50%) larger than the default value (“adjust” argument set to 1.5). This conservative choice aims to reduce the effect of the different sampling across c bins and to ensure that differences among distributions across c values are not the result of spurious sampling noise.Further, we transformed the kernel density function:$$begin{aligned} V(f_{dwell}) = -ln (p_{n}(f_{dwell})) end{aligned}$$
    (4)
    This (V(f_{dwell})) function, called potential (e.g.48), shows by design well-defined minima for the most frequently observed values of (f_{dwell}) (i.e. configurations most frequently observed for the community, which conform the modes of the probability distribution) in a given group of data. At these points, the potential exhibits a change of trend from decreasing to increasing, and therefore its derivative shows a change of sign. Eq. (4), thus, provides a simple criterion to identify possible system states, which is a reason why potentials have been used extensively across disciplines49,50,51. Nonetheless, because the position of extrema is invariant under the transformation, using probability distributions instead would not alter our conclusions.Representing the potential obtained from all the (f_{dwell}) system states associated with a same range of climatic driver c values allowed us to identify stable community configurations associated with a specific climate. The comparison of the potentials obtained for different c ranges enabled the description of how the community changed in response to climatic variation. The location of the minima revealed which states were stable for a given value of the climatic driver; the presence of two minima, then, flagged the existence of bistability (i.e. two different community compositions possible for the same c value).These minima are materialized as wells in the potential’s landscape, which provides an easy way to understand the concept of stability: the dynamics of the system for the given value of the driver will eventually “fall” into a well (either a state dominated by dwellers or a state dominated by tunnelers), with the shape of the well (e.g. its depth) determining how difficult it is for the system to “escape” that state. Therefore, the area inside a well provides quantification of the tendency of a system to stay in that specific state, i.e. the resilience of the associated ecological state or how strong a perturbation has to be to move the system from such an ecological state to another2,3,50,51,52,53. Thus, in addition to number and location of wells, measuring their associated area allowed us to further characterize the resilience of the community. To this end, we first set a visualization window common to all potentials. Specifically, we plotted the potentials within a range for the vertical variable (the potential, V) given by ([-1.5,1.5]); the horizontal variable (fraction of dwellers, (f_{dwell})) is by definition bounded between 0 and 1. For potentials that showed one single well, the area of the well was measured as the area above the potential curve within this visualization window. For potentials that showed two wells (bistability), we measured the value of the potential at the local maximum separating the two wells, and established that value as the upper (horizontal) line closing the area of each well. To ensure all cases were comparable and eliminate any arbitrariness of the choices above, we expressed resilience as a relative area; in other words, we further normalized the well area by the total area across wells for that potential, which means that any single-well case will show a resilience (or relative area) of 1, and the resilience of the two wells when there is bistability adds up to 1.Figure 1Left: Community composition by functional group for all weeks of observation1. Green represents dwellers, blue represents tunnelers, and orange represents the invader D. gazella. Right: Sketch of responses of the community composition to the climatic driver (i.e. phase diagram) expected from the physiological and behavioral characteristics of the functional groups in the community as described in text: linear (red), or non-linear but monotonic without (blue) or with (brown) hysteresis.Full size imageIdentifying ecological transitionsMeasuring a state variable, (f_{dwell}), and a driver, c (order and control parameter, respectively, in the jargon of regime shift theory), allowed us to study how their observed behavior over time materializes in a driver-state relationship (the so-called phase diagram) defining the possible shifts in dominance (i.e. regime shifts) that the community may undergo as climate changes12. The non-monotonic temporal behavior of the components of the order parameter (i.e. dwellers and tunneler availability) and the components of the control parameter (i.e. temperature and relative humidity) makes it difficult to predict the shape of the phase diagram, and therefore whether we can expect alternative stable states in the focal example. For such cases, the dominance of the dung beetle community could (1) shift in a linear fashion toward the functional group favored by climatic conditions; (2) shift between functional groups in non-linear threshold response to climatic conditions without hysteresis; or (3) shift between functional groups in non-linear threshold response to climatic conditions with hysteresis –and thus showing bistability (see Fig. 1b, or12). Other possibilities, e.g. a non-linear shift between functional groups where one group is favored at intermediate climatic conditions12 are discarded as the invader is better suited for warmer and drier conditions. To evaluate which of these possibilities occurred, we represented (f_{dwell}) as a function of c, as well as the location of the minima shown by the potentials above. In addition to the emerging shape of this relationship, this plot can reveal the presence of alternative stable states if two or more different points occur for the same value of the control parameter, c. More