Ecology

Department of Zoology, University of Cambridge, Cambridge, UKYucheng Wang, Bianca De Sanctis, Ruairidh Macleod, Daniel Money & Eske WillerslevLundbeck Foundation GeoGenetics Centre, Globe Institute, University of Copenhagen, Copenhagen, DenmarkYucheng Wang, Ana Prohaska, Jialu Cao, Antonio Fernandez-Guerra, James Haile, Kurt H. Kjær, Thorfinn Sand Korneliussen, Nicolaj Krog Larsen, Ruairidh Macleod, Hugh McColl, Mikkel Winther Pedersen, Fernando Racimo, Alexandra Rouillard, Anthony H. Ruter, Lasse Vinner, David J. Meltzer & Eske WillerslevALPHA, State Key Laboratory of Tibetan Plateau Earth System, Environment and Resources (TPESER), Institute of Tibetan Plateau Research (ITPCAS), Chinese Academy of Sciences (CAS), Beijing, ChinaYucheng WangKey Laboratory of Western China’s Environmental Systems (Ministry of Education), College of Earth and Environmental Science, Lanzhou University, Lanzhou, ChinaHaoran DongGénomique Métabolique, Genoscope, Institut François Jacob, CEA, CNRS, Univ Evry, Université Paris-Saclay, Evry, FranceAdriana Alberti, France Denoeud & Patrick WinckerInstitute for Integrative Biology of the Cell (I2BC), Université Paris-Saclay, CEA, CNRS, Gif-sur-Yvette, FranceAdriana AlbertiThe Arctic University Museum of Norway, UiT—The Arctic University of Norway, Tromsø, NorwayInger Greve Alsos, Eric Coissac, Galina Gusarova, Youri Lammers & Marie Kristine Føreid MerkelDepartment of Geography and Environment, University of Hawaii, Honolulu, HI, USADavid W. BeilmanDepartment of Geosciences and Natural Resource Management, University of Copenhagen, Copenhagen, DenmarkAnders A. BjørkInstitute of Earth Sciences, St Petersburg State University, St Petersburg, RussiaAnna A. Cherezova & Grigory B. FedorovArctic and Antarctic Research Institute, St Petersburg, RussiaAnna A. Cherezova & Grigory B. FedorovUniversité Grenoble-Alpes, Université Savoie Mont Blanc, CNRS, LECA, Grenoble, FranceEric CoissacDepartment of Genetics, University of Cambridge, Cambridge, UKBianca De Sanctis & Richard DurbinCarlsberg Research Laboratory, Copenhagen V, DenmarkChristoph Dockter & Birgitte SkadhaugeSchool of Geography and Environmental Science, University of Southampton, Southampton, UKMary E. EdwardsAlaska Quaternary Center, University of Alaska Fairbanks, Fairbanks, AK, USAMary E. EdwardsSchool of Environment, Earth and Ecosystem Sciences, The Open University, Milton Keynes, UKNeil R. Edwards & Philip B. HoldenCenter for the Environmental Management of Military Lands, Colorado State University, Fort Collins, CO, USAJulie EsdaleDepartment of Earth and Atmospheric Sciences, University of Alberta, Edmonton, Alberta, CanadaDuane G. FroeseFaculty of Biology, St Petersburg State University, St Petersburg, RussiaGalina GusarovaDepartment of Glaciology and Climate, Geological Survey of Denmark and Greenland, Copenhagen K, DenmarkKristian K. KjeldsenDepartment of Earth Science, University of Bergen, Bergen, NorwayJan Mangerud & John Inge SvendsenBjerknes Centre for Climate Research, Bergen, NorwayJan Mangerud & John Inge SvendsenDepartment of Geology, Quaternary Sciences, Lund University, Lund, SwedenPer MöllerCenter for Macroecology, Evolution and Climate, Globe Institute, University of Copenhagen, Copenhagen Ø, DenmarkDavid Nogués-Bravo, Hannah Lois Owens & Carsten RahbekCentre d’Anthropobiologie et de Génomique de Toulouse, Faculté de Médecine Purpane, Université Paul Sabatier, Toulouse, FranceLudovic OrlandoCenter for Global Mountain Biodiversity, Globe Institute, University of Copenhagen, Copenhagen, DenmarkHannah Lois Owens & Carsten RahbekGates of the Arctic National Park and Preserve, US National Park Service, Fairbanks, AK, USAJeffrey T. RasicDepartment of Geosciences, UiT—The Arctic University of Norway, Tromsø, NorwayAlexandra RouillardZoological Institute, Russian academy of sciences, St Petersburg, RussiaAlexei TikhonovResource and Environmental Research Center, Chinese Academy of Fishery Sciences, Beijing, ChinaYingchun XingCollege of Plant Science, Jilin University, Changchun, Jilin, ChinaYubin ZhangDepartment of Anthropology, Southern Methodist University, Dallas, TX, USADavid J. MeltzerWellcome Trust Sanger Institute, Wellcome Genome Campus, Cambridge, UKEske WillerslevMARUM, University of Bremen, Bremen, GermanyEske WillerslevAll authors contributed to the conception of the presented ideas. Y.W. and H.D. analysed the data. Y.W., D.J.M., A.P. and E.W. wrote the paper with inputs from all authors. More

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Time using the rock as refugeTemperature had an effect in the refuge usage of both species when analysed together (lme.zig: F3,192 = 7.97, p = 0.0001; Fig. 1A). However, species behaved differently (lme.zig: F1,192 = 14.79, p = 0.0004; Fig. 1A). As hypothesised, there was an interaction between temperature and species (lme.zig: F3,192 = 11.90, p  0.14, Fig. 1B).Size had an effect in the time exploring the rock (lme: F1,192 = 6.91, p = 0.012, Fig. 3) when species were analysed together, but there was no interaction with temperatures (lme: F3,192 = 0.42, p = 0.74, Fig. 3). We found that the interaction between species and size was close to be significant (lme: F1,192 = 3.62, p = 0.064, Fig. 3), implying that possibly smaller fish spent more time exploring the rock than bigger fish. However, when analysed separately, we did not find an effect of size in the exploring behaviour neither for twoline skiffias (lme: F1,96 = 2.99, p = 0.099, Fig. 3) nor for guppies (lme: F1,96 = 0.33, p = 0.569, Fig. 3).Figure 3Proportion of the total time observed (600 s) fish of different sizes spent exploring the rock. Lines represent the areas where the density of data is higher.Full size imageTime spent swimmingTemperature had an effect in the time spent swimming for both species when analysed together (lme: F3,192 = 23.48, p  More

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Area-based sub-targetWe found that 13.2% of Asian terrestrial landscapes were covered by PAs by the target date for Aichi 11 based on our in-country sources. However, it was 17.4% lower based on WDPA data (10.9%). The average increase in coverage across Asia during the 2010s was 0.4% ± SE 0.1% per year. PA coverage at the level of individual countries increased from a mean 11.1% in 2010 (SE = 1.4%) to 14.1% by 2020 (SE = 1.8%) based on our in-country sources, which was 16.5% higher than WDPA data (12.1 ± SE 1.6%). However, these overall figures concealed considerable country-level and sub-regional heterogeneity.A total of 8,673,433 km2 across 10 countries, equaling 19.6% of Asian terrestrial landscapes was managed as hunting concessions, governed by governments, communities or private sectors, but these areas have not been included in the countries’ report to the Protected Planet Initiative databases. Most of these areas are locally important in terms of biodiversity conservation and local socioeconomic outcomes which may qualify them as examples of “other effective area-based conservation measures” (OECMs). The increase in area-based conservation coverage represented by these areas, above the current Protected Planet Initiative statistic, ranged from 0.2% (Iran) to 41.4% (Russia). With that update incorporated, a total of 32.9% of Asian terrestrial landscapes are under protection, either as protected areas or hunting concessions (potentially as one type of OECMs).We found that 40% of Asian countries met a target of 17% coverage for PAs by 2020 based on our in-country sources, mainly in East and some South Asia, whereas West and Central Asian countries had generally not achieved this target (Figs. 1 and 2). We did not find any statistically significant association between the proportions of highly at-risk (CR/EN) mammalian species range outside PAs and the % PA extent in 2020 (β = −0.22 ± SE 0.15, t = −1.51, P = 0.14 in a Generalized Linear Model). The highest proportions of the highly at-risk (CR/EN) mammalian species range outside PAs were seen in West (βCR/EN_outsidePA = 1.77 ± SE 0.46, t = 3.86, P 10%, but Kuwait lost area. In East Asia, all countries showed at least some PA expansion (South Korea and Japan by >10%) whereas in Central Asia, almost no change was seen. It is also noteworthy that between 2010 and 2015, agricultural lands increased by 2.0% across the continent, averaging 0.51 ± SE 0.03% per year at country level, although 18 counties (45.0%) had agricultural land loss, mainly in West and Central Asia (12 out of 18 countries with agricultural land loss; Fig. 2).In our attempt to model the variation in achievement of area-based target (% PA extent), we found a single model with a ΔAICc weight of 1.0 (R2adj = 0.66; Table 1). There was no evidence to reject the null hypothesis that the model fits well (P = 0.99). This model included the predictors % agricultural extent in 2015, % PA extent in 2010, and sub-region (Table 1). Specifically, the coefficients suggested that countries with greater PA extent in 2010 and a smaller percentage of agricultural lands in 2015 were more likely to achieve higher percentage of PA extent by 2020 (βPAExtent2020 = 0.58 ± SE 0.10, t = 5.74, P  0.05).Table 2 Results of generalized linear models testing different hypotheses on the association between the percentage of ecoregions protected by the PA network in 2020 and ecological and geopolitical factors in Asian countries.Full size tableFor the coverage of highly at-risk (CR/EN) mammalian species, a single statistical model was also selected, with non-significant deviance goodness of fit (P = 0.83), which included only the % PA extent by 2020 and Region as predictors (R2adj = 0. 27). Although there was no evidence for association between the % PA extent by 2020 and the coverage of threatened species (βPAExtent2020 = −0.23 ± SE 0.15, t = −1.57, P = 0.13). However, the coverage of threatened species varied geographically, with high intercept differences for East Asia (βEastAsia = −0.23 ± SE 0.15, t = −1.57, P = 0.13), implying the largest median of range of highly at-risk (CR/EN) mammalian species outside the current network of PAs within each country.PA management effectiveness sub-targetFor the level of PAME assessment, we found that out of 22781 PAs within the 40 studied Asian countries, only 7.0% have been assessed based on PAME criteria (n = 1599), averaging 17.4% ± of PAs per country (SE = 2.5%). Israel, Japan, Lao, Bahrain, Oman and Qatar had no PA assessed based on the PAME criteria while over 1/3 of PAs in Indonesia, Cambodia, Bhutan, Jordan, Nepal, Turkey, Singapore and the UAE were PAME assessed. When modeling the level of PAME assessment, three best supported models were averaged (Table 3), with the averaged model including GDP2019, % PA extent 2020 and the Region as predictors. The averaged model coefficients would be non-significant under a hypothesis-testing approach (βGDP2019 = −0.18 ± SE 0.12, t = 1.47, P = 0.14 and βPAExtent2020 = −0.15 ± SE 0.11, t = 1.31, P = 0.19). Similarly, there was no evidence for the association between the ratio of PAs with PAME and Asian regions (P  > 0.05).Table 3 Results of generalized linear models testing different hypotheses on the association between the ratio of PAs with management effectiveness (PAME) in 2020 and ecological and geopolitical factors in Asian countries.Full size table More

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Differences in environmental factorsEnvironmental factors showed value differences between forest types, while the significance of differences differed among variables, which were both found with corrected values and original measurements (Fig. 1).Figure 1The differences in environmental factors between naturally regenerating forests (NF) and planted forests (PF) in China. The environmental factors include three annual climatic factors (a–c), three seasonal temperature factors (d–f), three seasonal precipitation factors (g–i), three biotic factors (j–l), and two soil factors (m,n). Three annual climatic factors include mean annual air temperature (MAT, a), mean annual precipitation (MAP, b), and aridity index (AI, c) defined as the ratio of MAP to annual potential evapotranspiration. Three seasonal temperature factors include the temperature of the warmest month (Tw, d), the temperature of the coldest month (Tc, e), temperature annual range (TR, f). Three seasonal precipitation factors include precipitation of the wettest month (Pw, g), precipitation of the driest month (Pd, h), and precipitation seasonality (Ps, i) defined as the standard deviation of monthly precipitation during the measuring year. Three biological factors include the mean annual leaf area index (LAI, j), the maximum leaf area index (MLAI, k), and stand age (SA, l). Two soil factors include soil organic carbon content (SOC, m) and soil total nitrogen content (STN, n). The differences are tested for each variable with one-way analysis of variance (ANOVA), where * and ** indicate significant differences between forest types at significance levels of α = 0.05 and α = 0.01, respectively. The corrected values are mean values during 2003–2019 after correcting the original measurements with the interannual trend (See methods), which are listed in each panel, while original measurements are mean values during the measuring period of each ecosystem, which are not shown in each panel.Full size imageFor annual climatic factors, the significant difference between NF and PF only appeared in MAT (Fig. 1a). The mean MAT of NF was 10.50 ± 7.81 °C, which was significantly lower than that of PF (15.65 ± 6.23 °C) (p  0.05) (Fig. 2c). Even considering the significant effects of MAT on ER, ANCOVA results obtained by fixing MAT as a covariant also suggested that ER values did not significantly differ between forest types (F = 0.01, p  > 0.05). Fixing other variables as a covariant also drew a similar result.Therefore, NF showed a lower NEP resulting from the lower GPP than PF, while their differences were not statistically significant (Fig. 2).Differences in NEP latitudinal patternsCarbon fluxes showed divergent latitudinal patterns between NF and PF, while their latitudinal patterns varied among carbon fluxes, which were both found with corrected values and original measurements (Fig. 3).Figure 3The latitudinal patterns of carbon fluxes over Chinese naturally regenerating forests (NF) and planted forests (PF). The carbon fluxes include net ecosystem productivity (NEP, a,b), gross primary productivity (GPP, c,d), and ecosystem respiration (ER, e,f). Each panel is drawn with the corrected values (blue points) and original measurements (grey points), respectively. The blue and black lines represent the regression lines calculated from the corrected values and original measurements, respectively, with their regression statistics listed in blue and black letters. Only the regression slope (Sl) and R2 of each regression are listed. The grey lines represent the regressions between carbon fluxes added by random errors and latitude. Only significant (p  0.05).The ER of NF showed a significant decreasing latitudinal pattern (Fig. 3e), while that of PF exhibited no significant latitudinal pattern (Fig. 3f). The increasing latitude caused the ER of NF to significantly decrease. Each unit increase in latitude led to a 28.71 gC m−2 year−1 decrease in ER, with an R2 of 0.31. However, the increasing latitude contributed little to the ER spatial variation of PF (p  > 0.05).In addition, the latitudinal patterns of carbon fluxes and their differences between forest types were also obtained with the original measurements (Fig. 3, grey points). The latitudinal patterns of random error adding carbon fluxes were comparable to those of our corrected carbon fluxes (Fig. 3), which confirmed that the latitudinal patterns of carbon fluxes and their differences between forest types would not be affected by the uncertainties in generating the corrected carbon fluxes.Therefore, among NFs, the similar decreasing latitudinal patterns of GPP and ER meant that NEP showed no significant latitudinal pattern, while the significant decreasing latitudinal pattern of GPP and no significant latitudinal pattern of ER caused NEP to show a decreasing latitudinal pattern among PFs.Differences in the environmental effects on NEP spatial variationsEnvironmental factors, including the annual climatic factors, seasonal temperature factors, seasonal precipitation factors, biological factors, and soil factors, exerted divergent effects on the spatial variations of NEP and its components, which also differed between forest types (Table 1). No factor was found to affect that the spatial variation of NEP among NFs, while most annual and seasonal climatic factors were found to affect that among PFs. The spatial variations of GPP and ER among NFs were both affected by most annual and seasonal climatic factors and LAI, while those among PFs were primarily shaped by most annual and seasonal climatic factors. Though LAI showed no significant effect on GPP and ER spatial variations among PFs, SA exerted a significant negative effect. In addition, the spatial variations of soil variables contributed little to the spatial variations of carbon fluxes. Therefore, among NFs, most annual and seasonal climatic factors and LAI were found to affect GPP and ER spatial variations, while no factor was found to significantly influent the NEP spatial variation. However, among PFs, most annual and seasonal climatic factors were found to affect the spatial variations of NEP and its components, while LAI showed no significant effect. Using the original measurements also generated the similar correlation coefficients (Supplementary Table S1).Table 1 Correlation coefficients between carbon fluxes and environmental factors in naturally regenerating forests (NF) and planted forests (PF).Full size tableGiven the high correlations among annual climatic factors and seasonal climatic factors (Supplementary Table S2), the partial correlation analysis was applied to determine which factors should be employed to reveal the mechanisms underlying the spatial variations of NEP. Partial correlation analysis showed that MAT and MAP exerted the most important roles in spatial variations of NEP and its components (Table 2). After controlling MAT (or MAP), other factors seldom showed significant correlation with carbon fluxes, especially fixing MAT (Table 2). In addition, MAT and MAP exerted similar effects on the spatial variations of NEP and its components (Table 1). Using the original measurements also generated the similar partial correlation coefficients (Supplementary Table S3). Therefore, we only presented the effects of MAT on carbon flux spatial variations and their differences between forest types in detail.Table 2 Partial correlation coefficients between carbon fluxes and environmental factors in naturally regenerating forests (NF) and planted forests (PF) with fixing mean annual air temperature (MAT) or mean annual precipitation (MAP).Full size tableThe increasing MAT increased carbon fluxes, while the increasing rates differed between forest types (Fig. 4). The increasing MAT contributed little to the NEP spatial variation of NF but raised the NEP of PF (Fig. 4a,b). Each unit increase in MAT caused the NEP of PF to increase at a rate of 27.77 gC m−2 year−1, with an R2 of 0.31 (Fig. 4b). The increasing MAT significantly raised GPP in NF and PF (Fig. 4c,d). For NF, each unit increase in MAT increased GPP at a rate of 43.76 gC m−2 year−1, with an R2 of 0.49 (Fig. 4c), while each unit increase in MAT increased the GPP of PF at a rate of 69.18 gC m−2 year−1, with an R2 of 0.57 (Fig. 4d). The GPP increasing rates did not significantly differ between NF and PF (F = 1.52, p  > 0.05). The increasing MAT also raised ER in both NF and PF (Fig. 4e,f), whose increasing rates were 38.97 gC m−2 year−1 (Fig. 4e) and 36.79 gC m−2 year−1 (Fig. 4f), respectively, while their differences were not statistically significant (F = 0.01, p  > 0.05). In addition, using the original measurements also generated the similar spatial variations and their differences between forest types (Fig. 4). Furthermore, the random error adding carbon fluxes responded similarly to those of our correcting carbon fluxes (Fig. 4), indicating that the effects of MAT on carbon fluxes would not be affected by the uncertainties in our correcting carbon fluxes. Therefore, the similar responses of GPP and ER to MAT made MAT contribute little to NEP spatial variations among NFs, while GPP and ER showed divergent response rates to MAT, which made NEP increase with MAT among PFs.Figure 4The effects of mean annual air temperature (MAT) on the spatial variations of carbon fluxes over Chinese naturally regenerating forests (NF) and planted forests (PF). The carbon fluxes include net ecosystem productivity (NEP, a,b), gross primary productivity (GPP, c,d), and ecosystem respiration (ER, e,f). Each panel is drawn with the corrected values (blue points) and original measurements (grey points), respectively. The blue and black lines represent the regression lines calculated from the corrected values and original measurements, respectively, with their regression statistics listed in blue and black letters. Only the regression slope (Sl) and R2 of each regression are listed. The grey lines represent the regressions between carbon fluxes added by random errors and latitude. Only significant (p  More

Alpha diversity differences among communitiesNematode gut microbiomes were assigned into their respective species categories of E. antarcticus and P. murrayi based on 18S host data that was consistent with morphology (see Methods “Microinvertebrate haplotypes”). In contrast, due to recovery of three undiscernible 18S tardigrade haplotypes, the gut microbiomes were assigned to Tardigrada. Mat bacterial communities were significantly (Tukey’s HSD, P  0.65, χ2(1)  0.38, χ2(3)  More