Ecology

Carothers, J. H. & Jaksić, F. M. Time as a Niche difference: The role of interference competition. Oikos 42, 403–406 (1984).
Google Scholar 
Kronfeld-Schor, N. & Dayan, T. Partitioning of time as an ecological resource. Annu. Rev. Ecol. Evol. Syst. 34, 153–181 (2003).
Google Scholar 
Lesmeister, D. B., Nielsen, C. K., Schauber, E. M. & Hellgren, E. C. Spatial and temporal structure of a mesocarnivore guild in midwestern North America. Wildl. Monogr. 191, 1–61 (2015).
Google Scholar 
Chmura, H. E. et al. Plasticity and repeatability of activity patterns in free-living Arctic ground squirrels. Anim. Behav. 169, 81–91 (2020).
Google Scholar 
Helm, B. et al. Two sides of a coin: Ecological and chronobiological perspectives of timing in the wild. Philos. Trans. R. Soc. B Biol. Sci. 372, 20160246 (2017).
Google Scholar 
Alós, J., Martorell-Barceló, M. & Campos-Candela, A. Repeatability of circadian behavioural variation revealed in free-ranging marine fish. R. Soc. Open Sci. 4, 160791 (2017).PubMed 
PubMed Central 

Google Scholar 
Schlicht, L. & Kempenaers, B. The effects of season, sex, age and weather on population-level variation in the timing of activity in Eurasian Blue Tits Cyanistes caeruleus. Ibis 162, 1146–1162 (2020).
Google Scholar 
Helm, B. & Visser, M. E. Heritable circadian period length in a wild bird population. Proc. R. Soc. B Biol. Sci. 277, 3335–3342 (2010).
Google Scholar 
Nikhil, K. L., Abhilash, L. & Sharma, V. K. Molecular correlates of circadian clocks in fruit fly drosophila melanogaster populations exhibiting early and late emergence chronotypes. J. Biol. Rhythms 31, 125–141 (2016).CAS 
PubMed 

Google Scholar 
Allebrandt, K. V. et al. CLOCK gene variants associate with sleep duration in two independent populations. Biol. Psychiatry 67, 1040–1047 (2010).CAS 
PubMed 

Google Scholar 
Maukonen, M. et al. Genetic associations of chronotype in the finnish general population. J. Biol. Rhythms 35, 501–511 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Roecklein, K. A. et al. Melanopsin gene variations interact with season to predict sleep onset and chronotype. Chronobiol. Int. 29, 1036–1047 (2012).CAS 
PubMed 
PubMed Central 

Google Scholar 
Steinmeyer, C., Kempenaers, B. & Mueller, J. C. Testing for associations between candidate genes for circadian rhythms and individual variation in sleep behaviour in blue tits. Genetica 140, 219–228 (2012).CAS 
PubMed 

Google Scholar 
Stuber, E. F., Baumgartner, C., Dingemanse, N. J., Kempenaers, B. & Mueller, J. C. Genetic correlates of individual differences in sleep behavior of free-living great tits (Parus major). G3 GenesGenomesGenetics 6, 599–607 (2016).CAS 

Google Scholar 
Cuthill, I. C. & Macdonald, W. A. Experimental manipulation of the dawn and dusk chorus in the blackbird Turdus merula. Behav. Ecol. Sociobiol. 26, 209–216 (1990).
Google Scholar 
Grava, T., Grava, A. & Otter, K. A. Supplemental feeding and dawn singing in black-capped chickadees. Condor 111, 560–564 (2009).
Google Scholar 
Saggese, K., Korner-Nievergelt, F., Slagsvold, T. & Amrhein, V. Wild bird feeding delays start of dawn singing in the great tit. Anim. Behav. 81, 361–365 (2011).
Google Scholar 
Dominoni, D. M. Effects of artificial light at night on daily and seasonal organization of European blackbirds (Turdus merula). https://kops.uni-konstanz.de/handle/123456789/32198 Accessed 23 February 2022 (2013).
Lehmann, M., Spoelstra, K., Visser, M. E. & Helm, B. Effects of temperature on circadian clock and chronotype: An experimental study on a passerine bird. Chronobiol. Int. 29, 1062–1071 (2012).PubMed 

Google Scholar 
Zsebők, S. et al. Short- and long-term repeatability and pseudo-repeatability of bird song: Sensitivity of signals to varying environments. Behav. Ecol. Sociobiol. 71, 154 (2017).
Google Scholar 
Raap, T., Pinxten, R. & Eens, M. Artificial light at night disrupts sleep in female great tits (Parus major) during the nestling period and is followed by a sleep rebound. Environ. Pollut. 215, 125–134 (2016).CAS 
PubMed 

Google Scholar 
Grunst, M. L., Grunst, A. S., Pinxten, R. & Eens, M. Variable and consistent traffic noise negatively affect the sleep behavior of a free-living songbird. Sci. Total Environ. 778, 146338 (2021).CAS 
PubMed 

Google Scholar 
Gaynor, K. M., Hojnowski, C. E., Carter, N. H. & Brashares, J. S. The influence of human disturbance on wildlife nocturnality. Science 360, 1232–1235 (2018).CAS 
PubMed 

Google Scholar 
Stuber, E. F. et al. Perceived predation risk affects sleep behaviour in free-living great tits Parus major. Anim. Behav. 98, 157–165 (2014).
Google Scholar 
Niemelä, P. T. & Dingemanse, N. J. Individual versus pseudo-repeatability in behaviour: Lessons from translocation experiments in a wild insect. J. Anim. Ecol. 86, 1033–1043 (2017).PubMed 

Google Scholar 
Garamszegi, L. Z. & Møller, A. P. Partitioning within-species variance in behaviour to within- and between-population components for understanding evolution. Ecol. Lett. 20, 599–608 (2017).PubMed 

Google Scholar 
Niemelä, P. T. & Dingemanse, N. J. On the usage of single measurements in behavioural ecology research on individual differences. Anim. Behav. 145, 99–105 (2018).
Google Scholar 
Browne, W. J., McCleery, R. H., Sheldon, B. C. & Pettifor, R. A. Using cross-classified multivariate mixed response models with application to life history traits in great tits (Parus major). Stat. Model. 7, 217–238 (2007).MathSciNet 
MATH 

Google Scholar 
Pettifor, R. A., Sheldon, B. C., Browne, W. J., Rasbash, J. & McCleery, R.
H. Partitioning of Phenotypic Variance in Life-history Traits in the Great Tit, Parus major.
https://seis.bristol.ac.uk/~frwjb/materials/phenovar.pdf (2003). Accessed 23 February 2022.Casasole, G. et al. Neither artificial light at night, anthropogenic noise nor distance from roads are associated with oxidative status of nestlings in an urban population of songbirds. Comp. Biochem. Physiol. A 210, 14–21 (2017).CAS 

Google Scholar 
Payevsky, V. A. Mortality rate and population density regulation in the great tit, Parus major L.: A review. Russ. J. Ecol. 37, 180 (2006).
Google Scholar 
Vermeulen, A., Eens, M., Van Dongen, S. & Müller, W. Does baseline innate immunity change with age? A multi-year study in great tits. Exp. Gerontol. 92, 67–73 (2017).CAS 
PubMed 

Google Scholar 
Haftorn, S. Incubation during the egg-laying period in relation to clutch-size and other aspects of reproduction in the great tit Parus major. Ornis Scand. Scand. J. Ornithol. 12, 169–185 (1981).
Google Scholar 
Grunst, M. L., Grunst, A. S., Pinxten, R., Eens, G. & Eens, M. An experimental approach to investigating effects of artificial light at night on free-ranging animals: Implementation, results and directions for future research. J. Vis. Exp. 180, e63381 (2022).

Google Scholar 
Halfwerk, W. et al. Low-frequency songs lose their potency in noisy urban conditions. Proc. Natl. Acad. Sci. 108, 14549–14554 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Specht, R. Avisoft-saslab pro: Sound analysis and synthesis laboratory. Avis. Bioacoustics
http://avisoft.com/SASLab_deutsch.pdf Accessed 23 February 2022 (2002).Iserbyt, A., Griffioen, M., Borremans, B., Eens, M. & Müller, W. How to quantify animal activity from radio-frequency identification (RFID) recordings. Ecol. Evol. 8, 10166–10174 (2018).PubMed 
PubMed Central 

Google Scholar 
Raap, T., Pinxten, R. & Eens, M. Light pollution disrupts sleep in free-living animals. Sci. Rep. 5, 13557 (2015).PubMed 
PubMed Central 

Google Scholar 
Meijdam, M., Müller, W., Thys, B. & Eens, M. No relationship between chronotype and timing of breeding when variation in daily activity patterns across the breeding season is taken into account. Ecol. Evol. 12, e9353 (2022).PubMed 
PubMed Central 

Google Scholar 
R Core Team. R: a language and environment for statistical computing. R Found. Stat. Comput. https://www.R-project.org/ Accessed 23 February 2022 (2013).Rousset, F. & Ferdy, J.-B. Testing environmental and genetic effects in the presence of spatial autocorrelation. Ecography 37, 781–790 (2014).
Google Scholar 
Araya-Ajoy, Y. G., Mathot, K. J. & Dingemanse, N. J. An approach to estimate short-term, long-term and reaction norm repeatability. Methods Ecol. Evol. 6, 1462–1473 (2015).
Google Scholar 
Mitchell, D. J., Dujon, A. M., Beckmann, C. & Biro, P. A. Temporal autocorrelation: A neglected factor in the study of behavioral repeatability and plasticity. Behav. Ecol. 31, 222–231 (2020).
Google Scholar 
Bell, A. M., Hankison, S. J. & Laskowski, K. L. The repeatability of behaviour: A meta-analysis. Anim. Behav. 77, 771–783 (2009).PubMed 
PubMed Central 

Google Scholar 
Graham, J. L., Cook, N. J., Needham, K. B., Hau, M. & Greives, T. J. Early to rise, early to breed: A role for daily rhythms in seasonal reproduction. Behav. Ecol. 28, 1266–1271 (2017).
Google Scholar 
Maury, C., Serota, M. W. & Williams, T. D. Plasticity in diurnal activity and temporal phenotype during parental care in European starlings Sturnus vulgaris. Anim. Behav. 159, 37–45 (2020).
Google Scholar 
Schlicht, L., Valcu, M., Loës, P., Girg, A. & Kempenaers, B. No relationship between female emergence time from the roosting place and extrapair paternity. Behav. Ecol. 25, 650–659 (2014).
Google Scholar 
Steinmeyer, C., Schielzeth, H., Mueller, J. C. & Kempenaers, B. Variation in sleep behaviour in free-living blue tits, Cyanistes caeruleus: Effects of sex, age and environment. Anim. Behav. 80, 853–864 (2010).
Google Scholar 
Stuber, E. F., Dingemanse, N. J., Kempenaers, B. & Mueller, J. C. Sources of intraspecific variation in sleep behaviour of wild great tits. Anim. Behav. 106, 201–221 (2015).
Google Scholar 
Raap, T., Pinxten, R. & Eens, M. Cavities shield birds from effects of artificial light at night on sleep. J. Exp. Zool. Part Ecol. Integr. Physiol. 329, 449–456 (2018).
Google Scholar 
Edelaar, P., Siepielski, A. M. & Clobert, J. Matching habitat choice causes directed gene flow: A neglected dimension in evolution and ecology. Evolution 62, 2462–2472 (2008).PubMed 

Google Scholar 
Gorissen, L. & Eens, M. Interactive communication between male and female great tits (Parus major) during the dawn chorus. Auk 121, 184–191 (2004).
Google Scholar 
Halfwerk, W., Bot, S. & Slabbekoorn, H. Male great tit song perch selection in response to noise-dependent female feedback. Funct. Ecol. 26, 1339–1347 (2012).
Google Scholar 
Steinmeyer, C., Mueller, J. C. & Kempenaers, B. Individual variation in sleep behaviour in blue tits Cyanistes caeruleus: Assortative mating and associations with fitness-related traits. J. Avian Biol. 44, 159–168 (2013).
Google Scholar 
Cain, J. R. & Wilson, W. O. The influence of specific environmental parameters on the circadian rhythms of chickens. Poult. Sci. 53, 1438–1447 (1974).CAS 
PubMed 

Google Scholar 
Zhang, Z. C. et al. Circadian clock genes are rhythmically expressed in specific segments of the hen oviduct. Poult. Sci. 95, 1653–1659 (2016).CAS 
PubMed 

Google Scholar 
Womack, R. J. Clocks in the wild: biological rhythms of great tits and the environment. https://theses.gla.ac.uk/81345/ Accessed 23 February 2022 (2020).Dominoni, D., Smit, J. A. H., Visser, M. E. & Halfwerk, W. Multisensory pollution: Artificial light at night and anthropogenic noise have interactive effects on activity patterns of great tits (Parus major). Environ. Pollut. 256, 113314 (2020).CAS 
PubMed 

Google Scholar 
Matthysen, E., Adriaensen, F. & Dhondt, A. A. Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus, Parus major). Glob. Change Biol. 17, 1–16 (2011).
Google Scholar  More

Hantson, S., Huxman, T. E., Kimball, S., Randerson, J. T. & Goulden, M. L. Warming as a driver of vegetation loss in the Sonoran Desert of California. J. Geophys. Res. Biogeosci. 126, e2020JG005942. https://doi.org/10.1029/2020JG005942 (2021).Article 
ADS 

Google Scholar 
Lortie, C. J., Filazzola, A., Kelsey, R., Hart, A. K. & Butterfield, H. S. Better late than never: A synthesis of strategic land retirement and restoration in California. Ecosphere 9, e02367. https://doi.org/10.1002/ecs2.2367 (2018).Article 

Google Scholar 
Ye, J.-S., Reynolds, J. F., Sun, G.-J. & Li, F.-M. Impacts of increased variability in precipitation and air temperature on net primary productivity of the Tibetan Plateau: A modeling analysis. Clim. Change 119, 321–332. https://doi.org/10.1007/s10584-013-0719-2 (2013).Article 
ADS 

Google Scholar 
Pendergrass, A. G., Knutti, R., Lehner, F., Deser, C. & Sanderson, B. M. Precipitation variability increases in a warmer climate. Sci. Rep. 7, 17966. https://doi.org/10.1038/s41598-017-17966-y (2017).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Zhang, W. et al. Increasing precipitation variability on daily-to-multiyear time scales in a warmer world. Sci. Adv. 7, eabf8021. https://doi.org/10.1126/sciadv.abf8021 (2021).Article 
ADS 
PubMed 
PubMed Central 

Google Scholar 
Stahle David, W. Anthropogenic megadrought. Science 368, 238–239. https://doi.org/10.1126/science.abb6902 (2020).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Williams, A. P. et al. Large contribution from anthropogenic warming to an emerging North American megadrought. Science 368, 314–318. https://doi.org/10.1126/science.aaz9600 (2020).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Bryant, B. P. et al. Shaping land use change and ecosystem restoration in a water-stressed agricultural landscape to achieve multiple benefits. Front. Sustain. Food Syst. 4, 138 (2020).Article 

Google Scholar 
Ross, C. W. et al. Woody-biomass projections and drivers of change in sub-Saharan Africa. Nat. Clim. Chang. 11, 449–455. https://doi.org/10.1038/s41558-021-01034-5 (2021).Article 
ADS 
PubMed 
PubMed Central 

Google Scholar 
Scanlon, B. R., Reedy, R. C., Stonestrom, D. A., Prudic, D. E. & Dennehy, K. F. Impact of land use and land cover change on groundwater recharge and quality in the southwestern US. Glob. Change Biol. 11, 1577–1593. https://doi.org/10.1111/j.1365-2486.2005.01026.x (2005).Article 
ADS 

Google Scholar 
Scanlon, B. R. et al. Global synthesis of groundwater recharge in semiarid and arid regions. Hydrol. Process. 20, 3335–3370. https://doi.org/10.1002/hyp.6335 (2006).Article 
ADS 
CAS 

Google Scholar 
Kelsey, R., Hart, A., Butterfield, H. S. & Vink, D. Groundwater sustainability in the San Joaquin Valley: Multiple benefits if agricultural lands are retired and restored strategically. Calif. Agric. 2, 151–154 (2018).Article 

Google Scholar 
Capdevila, P. et al. Reconciling resilience across ecological systems, species and subdisciplines. J. Ecol. 109, 3102–3113. https://doi.org/10.1111/1365-2745.13775 (2021).Article 

Google Scholar 
Thebault, A., Mariotte, P., Lortie, C. & MacDougall, A. Land management trumps the effects of climate change and elevated CO2 on grassland functioning. J. Ecol. 102, 896–904. https://doi.org/10.1111/1365-2745.12236 (2014).Article 

Google Scholar 
Turney, C., Ausseil, A.-G. & Broadhurst, L. Urgent need for an integrated policy framework for biodiversity loss and climate change. Nature Ecol. Evol. 4, 996–996. https://doi.org/10.1038/s41559-020-1242-2 (2020).Article 

Google Scholar 
Strassburg, B. B. N. et al. Global priority areas for ecosystem restoration. Nature 586, 724–729. https://doi.org/10.1038/s41586-020-2784-9 (2020).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Ellison, A. M. Foundation species, non-trophic interactions, and the value of being common. iScience 13, 254–268. https://doi.org/10.1016/j.isci.2019.02.020 (2019).Article 
ADS 
PubMed 
PubMed Central 

Google Scholar 
O’Brien, M. J., Carbonell, E. P., Losapio, G., Schlüter, P. M. & Schöb, C. Foundation species promote local adaptation and fine-scale distribution of herbaceous plants. J. Ecol. 109, 191–203. https://doi.org/10.1111/1365-2745.13461 (2021).Article 
CAS 

Google Scholar 
Bagley, J. E. et al. The influence of land cover on surface energy partitioning and evaporative fraction regimes in the U.S. Southern Great Plains. J. Geophys. Res.: Atmos. 122, 5793–5807. https://doi.org/10.1002/2017JD026740 (2017).Article 
ADS 

Google Scholar 
Norris, C., Hobson, P. & Ibisch, P. L. Microclimate and vegetation function as indicators of forest thermodynamic efficiency. J. Appl. Ecol. 49, 562–570. https://doi.org/10.1111/j.1365-2664.2011.02084.x (2012).Article 

Google Scholar 
Brooker, R. W. et al. Tiny niches and translocations: The challenge of identifying suitable recipient sites for small and immobile species. J. Appl. Ecol. 55, 621–630. https://doi.org/10.1111/1365-2664.13008 (2018).Article 

Google Scholar 
Forzieri, G. et al. Increased control of vegetation on global terrestrial energy fluxes. Nat. Clim. Chang. 10, 356–362. https://doi.org/10.1038/s41558-020-0717-0 (2020).Article 
ADS 

Google Scholar 
Milling, C. R. et al. Habitat structure modifies microclimate: An approach for mapping fine-scale thermal refuge. Methods Ecol. Evol. 9, 1648–1657. https://doi.org/10.1111/2041-210X.13008 (2018).Article 

Google Scholar 
Ghazian, N., Zuliani, M. & Lortie, C. J. Micro-climatic amelioration in a california desert: Artificial shelter versus shrub canopy. J. Ecol. Eng. 21, 216–228. https://doi.org/10.12911/22998993/126875 (2020).Article 

Google Scholar 
Wright, A. J., Barry, K. E., Lortie, C. J. & Callaway, R. M. Biodiversity and ecosystem functioning: Have our experiments and indices been underestimating the role of facilitation?. J. Ecol. 109, 1962–1968. https://doi.org/10.1111/1365-2745.13665 (2021).Article 

Google Scholar 
Germano, D. J. et al. The San Joaquin Desert of California: Ecologically misunderstood and overlooked. Nat. Areas J. 31, 138–147. https://doi.org/10.3375/043.031.0206 (2011).Article 

Google Scholar 
Fairbairn, M., LaChance, J., De Master, K. T. & Ashwood, L. In vino veritas, in aqua lucrum: Farmland investment, environmental uncertainty, and groundwater access in California’s Cuyama Valley. Agric. Hum. Values 38, 285–299. https://doi.org/10.1007/s10460-020-10157-y (2021).Article 

Google Scholar 
Filazzola, A., Lortie, C. J., Westphal, M. F. & Michalet, R. Species-specificity challenges the predictability of facilitation along a regional desert gradient. J. Veg. Sci. 1, 1–12. https://doi.org/10.1111/jvs.12909 (2020).Article 

Google Scholar 
Cutlar, H. C. Monograph of the North American species of the genus Ephedra. Ann. Mo. Bot. Gard. 26, 373–428 (1939).Article 

Google Scholar 
Hollander, J. L., Wall, S. B. V. & Baguley, J. G. Evolution of seed dispersal in North American Ephedra. Evol. Ecol. 24, 333–345. https://doi.org/10.1007/s10682-009-9309-1 (2010).Article 

Google Scholar 
Filazzola, A., Brown, C., Westphal, M. & Lortie, C. J. Establishment of a desert foundation species is limited by exotic plants and light but not herbivory or water. Appl. Veg. Sci. 1, 1–12. https://doi.org/10.1111/avsc.12515 (2020).Article 

Google Scholar 
Lortie, C. J., Gruber, E., Filazzola, A., Noble, T. & Westphal, M. The Groot effect: Plant facilitation and desert shrub regrowth following extensive damage. Ecol. Evol. 8, 706–715. https://doi.org/10.1002/ece3.3671 (2018).Article 
PubMed 

Google Scholar 
Lortie, C. J. et al. Telemetry of the lizard species Gambelia sila at Carrizo plain national monument. Figshare. Dataset. https://doi.org/10.6084/m9.figshare.8239667.v2 (2019).Article 

Google Scholar 
Braun, J., Westphal, M. & Lortie, C. J. The shrub Ephedra californica facilitates arthropod communities along a regional desert climatic gradient. Ecosphere 12, e03760. https://doi.org/10.1002/ecs2.3760 (2021).Article 

Google Scholar 
Terando, A., Youngsteadt, E., Meineke, E. & Prado, S. Accurate near surface air temperature measurements are necessary to gauge large-scale ecological responses to global climate change. Ecol. Evol. 8, 5233–5234. https://doi.org/10.1002/ece3.3972 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Tielborger, K. & Kadmon, R. Indirect effects in a desert plant community: Is competition among annuals more intense under shrub canopies?. Plant Ecol. 150, 53–63 (2000).Article 

Google Scholar 
Holzapfel, C., Tielbörger, K., Parag, H. A., Kigel, J. & Sternberg, M. Annual plant–shrub interactions along an aridity gradient. Basic Appl. Ecol. 7, 268–279. https://doi.org/10.1016/j.baae.2005.08.003 (2006).Article 

Google Scholar 
Jankju, M. Role of nurse shrubs in restoration of an arid rangeland: Effects of microclimate on grass establishment. J. Arid Environ. 89, 103–109. https://doi.org/10.1016/j.jaridenv.2012.09.008 (2013).Article 
ADS 

Google Scholar 
Baldelomar, M., Atala, C. & Molina-Montenegro, M. A. Top-down and Bottom-up effects deployed by a nurse shrub allow facilitating an endemic mediterranean orchid. Front. Ecol. Evol. 7, 466 (2019).Article 

Google Scholar 
Tielborger, K. & Kadmon, R. Temporal environmental variation tips the balance between facilitation and interference in desert plants. Ecology 81, 1544–1553. https://doi.org/10.1890/0012-9658(2000)081[1544:TEVTTB]2.0.CO;2 (2000).Article 

Google Scholar 
Walter, J. Effects of changes in soil moisture and precipitation patterns on plant-mediated biotic interactions in terrestrial ecosystems. Plant Ecol. https://doi.org/10.1007/s11258-018-0893-4 (2018).Article 

Google Scholar 
Schob, C., Armas, C. & Pugnaire, F. Direct and indirect interactions co-determine species composition in nurse plant systems. Oikos 122, 1371–1379. https://doi.org/10.1111/j.1600-0706.2013.00390.x (2013).Article 

Google Scholar 
Eldridge, D. J., Beecham, G. & Grace, J. B. Do shrubs reduce the adverse effects of grazing on soil properties?. Ecohydrology 8, 1503–1513. https://doi.org/10.1002/eco.1600 (2015).Article 

Google Scholar 
Nerlekar, A. N. & Veldman, J. W. High plant diversity and slow assembly of old-growth grasslands. Proc. Natl. Acad. Sci. 117, 18550. https://doi.org/10.1073/pnas.1922266117 (2020).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Tielbörger, K. et al. Middle-Eastern plant communities tolerate 9 years of drought in a multi-site climate manipulation experiment. Nat. Commun. 5, 5102. https://doi.org/10.1038/ncomms6102 (2014).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Griffin, D. & Anchukaitis, K. J. How unusual is the 2012–2014 California drought?. Geophys. Res. Lett. 41, 9017–9023. https://doi.org/10.1002/2014GL062433 (2014).Article 
ADS 

Google Scholar 
Data, U. C. In US Climate Data Product, New Cuyama, vol. 1. https://www.usclimatedata.com (2021).Gherardi, L. A. & Sala, O. E. Effect of interannual precipitation variability on dryland productivity: A global synthesis. Glob. Change Biol. 25, 269–276. https://doi.org/10.1111/gcb.14480 (2019).Article 
ADS 

Google Scholar 
Ding, Y., Li, Z. & Peng, S. Global analysis of time-lag and -accumulation effects of climate on vegetation growth. Int. J. Appl. Earth Obs. Geoinf. 92, 102179. https://doi.org/10.1016/j.jag.2020.102179 (2020).Article 

Google Scholar 
Liu, H. et al. Analysis of the time-lag effects of climate factors on grassland productivity in Inner Mongolia. Glob. Ecol. Conserv. 30, e01751. https://doi.org/10.1016/j.gecco.2021.e01751 (2021).Article 

Google Scholar 
Liancourt, P., Song, X., Macek, M., Santrucek, J. & Dolezal, J. Plant’s-eye view of temperature governs elevational distributions. Glob. Change Biol. 26, 4094–4103. https://doi.org/10.1111/gcb.15129 (2020).Article 
ADS 

Google Scholar 
Ryan, M. J. et al. Too dry for lizards: Short-term rainfall influence on lizard microhabitat use in an experimental rainfall manipulation within a pinon-juniper woodland. Funct. Ecol. https://doi.org/10.1111/1365-2435.12595 (2015).Article 

Google Scholar 
Moore, D., Stow, A. & Kearney, M. R. Under the weather?—The direct effects of climate warming on a threatened desert lizard are mediated by their activity phase and burrow system. J. Anim. Ecol. 87, 660–671. https://doi.org/10.1111/1365-2656.12812 (2018).Article 
PubMed 

Google Scholar 
Gaudenti, N., Nix, E., Maier, P., Westphal, M. F. & Taylor, E. N. Habitat heterogeneity affects the thermal ecology of an endangered lizard. Ecol. Evol. 11, 14843–14856. https://doi.org/10.1002/ece3.8170 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Lortie, C. J., Filazzola, A. & Sotomayor, D. A. Functional assessment of animal interactions with shrub-facilitation complexes: A formal synthesis and conceptual framework. Funct. Ecol. 30, 41–51. https://doi.org/10.1111/1365-2435.12530 (2016).Article 

Google Scholar 
Lortie, C. J. et al. Shrub and vegetation cover predict resource selection use by an endangered species of desert lizard. Sci. Rep. 10, 4884. https://doi.org/10.1038/s41598-020-61880-9 (2020).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
De Frenne, P. et al. Global buffering of temperatures under forest canopies. Nature Ecol. Evol. 3, 744–749. https://doi.org/10.1038/s41559-019-0842-1 (2019).Article 

Google Scholar 
Avolio, M. L. et al. Determinants of community compositional change are equally affected by global change. Ecol. Lett. 24, 1892–1904. https://doi.org/10.1111/ele.13824 (2021).Article 
PubMed 

Google Scholar 
Cook-Patton, S. C. et al. Protect, manage and then restore lands for climate mitigation. Nat. Clim. Chang. 11, 1027–1034. https://doi.org/10.1038/s41558-021-01198-0 (2021).Article 
ADS 

Google Scholar 
Hedden-Nicely, D. R. Climate change and the future of western US water governance. Nat. Clim. Chang. https://doi.org/10.1038/s41558-021-01141-3 (2021).Article 

Google Scholar 
Suggitt, A. J. et al. Extinction risk from climate change is reduced by microclimatic buffering. Nat. Clim. Chang. 8, 713–717. https://doi.org/10.1038/s41558-018-0231-9 (2018).Article 
ADS 

Google Scholar 
Hanson, R. T., Flint, L. E., Faunt, C. C., Gibbs, D. R. & Schmid, W. Hydrologic models and analysis of water availability in Cuyama Valley, California. In U.S. Geological Survey Scientific Investigations Report, 2015 1–126 (2015).John, S. In Encyclopedia of World Climatology (ed John, E. O.) 89–94 (Springer Netherlands, 2005).James-Jeremy, J. et al. A systems approach to restoring degraded drylands. J. Appl. Ecol. 50, 730–739. https://doi.org/10.1111/1365-2664.12090 (2013).Article 

Google Scholar 
Upson, J. E. & Worts, G. F. In Ground water in the Cuyama Valley, California. Report No. 1110B 1–82 (1951).Hanson, M. T., Randall, T. & Sweetkind, D. Cuyama Valley, California hydrologic study—an assessment of water availability. In U.S. Geological Survey Scientific Investigations Report 2014 1–4. https://doi.org/10.3133/fs20143075 (2014).Greicius, T. NASA data show California’s San Joaquin Valley Still Sinking. JPL 28, 1–9 (2017).
Google Scholar 
Döll, P. et al. Impact of water withdrawals from groundwater and surface water on continental water storage variations. J. Geodyn. 59–60, 143–156. https://doi.org/10.1016/j.jog.2011.05.001 (2012).Article 

Google Scholar 
Lortie, C. J. & Filazzola, A. US climate data, New Cuyama, CA, 2016–2017. Figshare 1, 2016–2017. https://doi.org/10.6084/m9.figshare.17162600.v1 (2021).Article 

Google Scholar 
Lortie, C. J. & Filazzola, A. Vegetation surveys in Cuyama Valley, CA, USA in 2016 and 2017 at the peak of megadrought. Knowl. Netw. Biocompl. 1, 1–15. https://doi.org/10.5063/F1MG7MZH (2021).Article 

Google Scholar 
Hickman, J. C. The Jepson Manual (University of California Press, 1996).
Google Scholar 
Villanueva-Almanza, L. & Fonseca, R. M. In Taxonomic review and geographic distribution of Ephedra (Ephedraceae) in Mexico. ACTA BOTANICA MEXICANA 96 (2011).Alfieri, F. J. & Mottola, P. M. Seasonal changes in the phloem of Ephedra californica Wats. Bot. Gaz. 144, 240–246 (1983).Article 

Google Scholar 
Hoffman, O., de-Falco, N., Yizhaq, H. & Boeken, B. Annual plant diversity decreases across scales following widespread ecosystem engineer shrub mortality. J. Veg. Sci. https://doi.org/10.1111/jvs.12372 (2016).Article 

Google Scholar 
Ivey, K. N. et al. Thermal ecology of the federally endangered blunt-nosed leopard lizard (Gambelia sila). Conserv. Physiol. 2020, 8. https://doi.org/10.1093/conphys/coaa014 (2020).Article 

Google Scholar 
Grimes, A. J., Corrigan, G., Germano, D. J. & Smith, P. T. Mitochondrial phylogeography of the endangered blunt-nosed leopard lizard, Gambelia sila. Southwestern Natural. 59, 38–46. https://doi.org/10.1894/F06-GC-233.1 (2014).Article 

Google Scholar 
Stewart, J. A. E. et al. Habitat restoration opportunities, climatic niche contraction, and conservation biogeography in California’s San Joaquin Desert. PLoS ONE 14, e0210766. https://doi.org/10.1371/journal.pone.0210766 (2019).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Germano, D. J., Rathbun, G. B. & Saslaw, L. R. Effects of grazing and invasive grasses on desert vertebrates in California. J. Wildl. Manag. 76, 670–682. https://doi.org/10.1002/jwmg.316 (2012).Article 

Google Scholar 
Moss, B. The water framework directive: Total environment or political compromise?. Sci. Total Environ. 400, 32–41. https://doi.org/10.1016/j.scitotenv.2008.04.029 (2008).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Denevan, W. M. The “Pristine Myth ” revisited. Geogr. Rev. 101, 576–591. https://doi.org/10.1111/j.1931-0846.2011.00118.x (2011).Article 

Google Scholar 
da Cunha, A. R. Evaluation of measurement errors of temperature and relative humidity from HOBO data logger under different conditions of exposure to solar radiation. Environ. Monit. Assess. 187, 236. https://doi.org/10.1007/s10661-015-4458-x (2015).Article 
PubMed 

Google Scholar 
Terando, A. J., Youngsteadt, E., Meineke, E. K. & Prado, S. G. Ad hoc instrumentation methods in ecological studies produce highly biased temperature measurements. Ecol. Evol. 7, 9890–9904. https://doi.org/10.1002/ece3.3499 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
Nature, I. I. U. f. C. o. The IUCN red list of threatened species. IUCN 2019-1 1–142 (2019).Lortie, C. J., Filazzola, A., Butterfield, H. S. & Westphal, M. Cuyama Micronet. Figshare 1, 1–6. https://doi.org/10.6084/m9.figshare.11888199.v2 (2020).Article 

Google Scholar 
Team, R. C. R: A Language and Environment for Statistical Computing. Vol. 4.2.1 (R foundation for Statistical Computing, 2022).Pinheiro, J., Bates, D., DebRoy, S. & Deepayan, S. nlme: Linear and nonlinear mixed effects models. CRAN 3, 1–153 (2021).
Google Scholar 
Pebesma, E. spacetime: Spatio-temporal data in R. J. Stat. Softw. 1(7), 2012. https://doi.org/10.18637/jss.v051.i07 (2012).Article 

Google Scholar 
Bates, D. et al. lme4: Linear mixed-effects models using “Eigen” and S4. CRAN 2020, 1–122 (2020).
Google Scholar 
Lenth, R. V. emmeans: Estimated marginal means. CRAN 1, 1–89 (2022).
Google Scholar  More

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Lal, R. Soil carbon sequestration impacts on global climate change and food security. Science 304, 1623–1627 (2004).ADS 
PubMed 

Google Scholar 
Torn, M. S., Vitousek, P. M. & Trumbore, S. E. The influence of nutrient availability on soil organic matter turnover estimated by incubations and radiocarbon modeling. Ecosystems 8, 352–372 (2005).
Google Scholar 
Schimel, J. P. & Schaeffer, S. M. Microbial control over carbon cycling in soil. Front. Microbiol. 3, 348 (2012).PubMed 
PubMed Central 

Google Scholar 
Ding, X., Liang, C., Zhang, B., Yuan, Y. & Han, X. Higher rates of manure application lead to greater accumulation of both fungal and bacterial residues in macroaggregates of a clay soil. Soil Biol. Biochem. 84, 137–146 (2015).
Google Scholar 
Liang, C., Schimel, J. P. & Jastrow, J. D. The importance of anabolism in microbial control over soil carbon storage. Nat. Microbiol. 2, 17105 (2017).PubMed 

Google Scholar 
Kögel-Knabner, I. The macromolecular organic composition of plant and microbial residues as inputs to soil organic matter: fourteen years on. Soil Biol. Biochem. 105, A3–A8 (2017).
Google Scholar 
Miltner, A., Bombach, P., Schmidt-Brücken, B. & Kästner, M. SOM genesis: microbial biomass as a significant source. Biogeochemistry 111, 41–55 (2012).
Google Scholar 
Cotrufo, M. F., Wallenstein, M. D., Boot, C. M., Denef, K. & Paul, E. The Microbial Efficiency-Matrix Stabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: Do labile plant inputs form stable soil organic matter?. Global Change Biol. 19, 988–995 (2013).ADS 

Google Scholar 
Sokol, N. W. & Bradford, M. A. Microbial formation of stable soil carbon is more efficient from belowground than aboveground input. Nat. Geosci. 12, 46–53 (2019).ADS 

Google Scholar 
Six, J., Frey, S. D., Thiet, R. K. & Batten, K. M. Bacterial and fungal contributions to carbon sequestration in agroecosystems. Soil Sci. Soc. Am. J. 70, 555–569 (2006).ADS 

Google Scholar 
Simpson, A. J., Simpson, M. J., Smith, E. & Kelleher, B. P. Microbially derived inputs to soil organic matter: Are current estimates too low?. Environ. Sci. Technol. 41, 8070–8076 (2007).ADS 
PubMed 

Google Scholar 
Liang, C., Fujinuma, R. & Balser, T. C. Comparing PLFA and amino sugars for microbial analysis in an Upper Michigan old growth forest. Soil Biol. Biochem. 40, 2063–2065 (2008).
Google Scholar 
Shao, P., Liang, C., Lynch, L., Xie, H. & Bao, X. Reforestation accelerates soil organic carbon accumulation: Evidence from microbial biomarkers. Soil Biol. Biochem. 131, 182–190 (2019).
Google Scholar 
Ma, S. et al. Effects of seven-year nitrogen and phosphorus additions on soil microbial community structures and residues in a tropical forest in Hainan Island, China. Geoderma 361, 114034 (2020).ADS 

Google Scholar 
Zelles, L. Fatty acid patterns of phospholipids and lipopolysaccharides in the characterisation of microbial communities in soil: A review. Biol. Fertil. Soils 29, 111–129 (1999).
Google Scholar 
Kong, A. Y. Y. et al. Microbial community composition and carbon cycling within soil microenvironments of conventional, low-input, and organic cropping systems. Soil Biol. Biochem. 43, 20–30 (2011).PubMed 
PubMed Central 

Google Scholar 
Müller, K., Marhan, S., Kandeler, E. & Poll, C. Carbon flow from litter through soil microorganisms: From incorporation rates to mean residence times in bacteria and fungi. Soil Biol. Biochem. 115, 187–196 (2017).
Google Scholar 
Amelung, W. Syntax of Referencing in Assessment Methods for Soil Carbon (Lewis Publishers, 2001).
Google Scholar 
Joergensen, R. & Wichern, F. Quantitative assessment of the fungal contribution to microbial tissue in soil. Soil Biol. Biochem. 40, 2977–2991 (2008).
Google Scholar 
Joergensen, R. G. Amino sugars as specific indices for fungal and bacterial residues in soil. Biol. Fert. Soils 54, 559–568 (2018).
Google Scholar 
Wang, X. et al. Distinct regulation of microbial processes in the immobilization of labile carbon in different soils. Soil Biol. Biochem. 142, 107723 (2020).
Google Scholar 
Wang, J., Chapman, S. J. & Yao, H. Incorporation of 13C-labelled rice rhizodeposition into soil microbial communities under different fertilizer applications. Appl. Soil Ecol. 101, 11–19 (2016).ADS 

Google Scholar 
Cui, S. et al. Long-term fertilization management affects the C utilization from crop residues by the soil micro-food web. Plant Soil 429, 335–348 (2018).
Google Scholar 
Liu, X., Zhang, X. & Herbert, S. Feeding China’s growing needs for grain. Nature 465, 420 (2010).ADS 
PubMed 

Google Scholar 
Edmeades, D. C. The long-term effects of manures and fertilisers on soil productivity and quality: A review. Nutr. Cycl. Agroecosys. 66, 165–180 (2003).
Google Scholar 
Chaparro, J., Sheflin, A., Manter, D. & Vivanco, J. Manipulating the soil microbiome to increase soil health and plant fertility. Biol. Fertil. Soils 48, 489–499 (2012).
Google Scholar 
Jin, X. et al. Enhanced conversion of newly-added maize straw to soil microbial biomass C under plastic film mulching and organic manure management. Geoderma 313, 154–162 (2018).ADS 

Google Scholar 
Chen, X., Li, Z., Liu, M., Jiang, C. & Che, Y. Microbial community and functional diversity associated with different aggregate fractions of a paddy soil fertilized with organic manure and/or NPK fertilizer for 20 years. J. Soil Sediment. 15, 292–301 (2014).
Google Scholar 
Wang, Y. et al. Soil aggregation regulates distributions of carbon, microbial community and enzyme activities after 23-year manure amendment. Appl. Soil Ecol. 111, 65–72 (2017).
Google Scholar 
Joergensen, R. G., Mäder, P. & Fließbach, A. Long-term effects of organic farming on fungal and bacterial residues in relation to microbial energy metabolism. Biol. Fert. Soils 46, 303–307 (2010).
Google Scholar 
Sun, H. et al. Soil microbial community and microbial residues respond positively to minimum tillage under organic farming in Southern Germany. Appl. Soil Ecol. 108, 16–24 (2016).
Google Scholar 
Heijboer, A. et al. Plant biomass, soil microbial community structure and nitrogen cycling under different organic amendment regimes; A 15N tracer-based approach. Appl. Soil Ecol. 107, 251–260 (2016).
Google Scholar 
Six, J., Elliott, E. T. & Paustian, K. Soil macroaggregate turnover and microaggregate formation: A mechanism for C sequestration under no-tillage agriculture. Soil Biol. Biochem. 32, 2099–2103 (2000).
Google Scholar 
Wall, D. et al. Soil Ecology and Ecosystem Services (Oxford University Press, 2012).
Google Scholar 
Helgason, B. L., Walley, F. L. & Germida, J. J. No-till soil management increases microbial biomass and alters community profiles in soil aggregates. Appl. Soil Ecol. 46, 390–397 (2010).
Google Scholar 
Blaud, A. et al. Dynamics of bacterial communities in relation to soil aggregate formation during the decomposition of 13C-labelled rice straw. Appl. Soil Ecol. 53, 1–9 (2012).
Google Scholar 
Tisdall, J. M. & Oades, J. M. Organic matter and water stable aggregates in soils. Eur. J. Soil Sci. 33, 141–163 (1982).
Google Scholar 
Bronick, C. J. & Lal, R. Soil structure and management: A review. Geoderma 124, 3–22 (2005).ADS 

Google Scholar 
Li, N. et al. Separation of soil microbial community structure by aggregate size to a large extent under agricultural practices during early pedogenesis of a Mollisol. Appl. Soil Ecol. 88, 9–20 (2015).
Google Scholar 
Bidisha, M., Joerg, R. & Yakov, K. Effects of aggregation processes on distribution of aggregate size fractions and organic C content of a long-term fertilized soil. Eur. J. Soil Biol. 46, 365–370 (2010).
Google Scholar 
Xiang, X. et al. Divergence in fungal abundance and community structure between soils under long-term mineral and organic fertilization. Soil Till. Res. 196, 104491 (2020).
Google Scholar 
Jin, X. et al. Long-term plastic film mulching and fertilization treatments changed the annual distribution of residual maize straw C in soil aggregates under field conditions: Characterization by 13C tracing. J. Soils Sediment. 18, 169–178 (2018).
Google Scholar 
Kemper, W. & Rosenau, R. Syntax of referencing. In Methods of Soil Analysis (ed. Klute, A.) (ASA and SSSA, 1986).
Google Scholar 
Bossio, D. A. & Scow, K. M. Impact of carbon and flooding on the metabolic diversity of microbial communities in soils. Appl. Environ. Microbiol. 61, 4043–4050 (1995).ADS 
PubMed 
PubMed Central 

Google Scholar 
Denef, K. et al. Community shifts and carbon translocation within metabolically-active rhizosphere microorganisms in grasslands under elevated CO2. Biogeosciences 4, 769–779 (2007).ADS 

Google Scholar 
Tavi, N. M. et al. Linking microbial community structure and allocation of plant-derived carbon in an organic agricultural soil using 13CO2 pulse-chase labelling combined with 13C-PLFA profiling. Soil Biol. Biochem. 58, 207–215 (2013).
Google Scholar 
Bach, E. M., Baer, S. G., Meyer, C. K. & Six, J. Soil texture affects soil microbial and structural recovery during grassland restoration. Soil Biol. Biochem. 42, 2182–2191 (2010).
Google Scholar 
Pan, F., Li, Y., Chapman, S. J., Khan, S. & Yao, H. Microbial utilization of rice straw and its derived biochar in a paddy soil. Sci. Total Environ. 559, 15–23 (2016).ADS 
PubMed 

Google Scholar 
Olsson, P. A. Signature fatty acids provide tools for determination of the distribution and interactions of mycorrhizal fungi in soil. FEMS Microbial Ecol. 29, 303–310 (1999).
Google Scholar 
Zhang, X. & Amelung, W. Gas Chromatographic determination of muramic acid, glucosamine, mannosamine, and galactosamine in soils. Soil Biol. Biochem. 28, 1201–1206 (1996).
Google Scholar 
Zhang, X. et al. Land-use effects on amino sugars in particle size fractions of an Argiudoll. Appl. Soil Ecol. 11, 271–275 (1999).
Google Scholar 
van Groenigen, K.-J. et al. Abundance, production and stabilization of microbial biomass under conventional and reduced tillage. Soil Biol. Biochem. 42, 48–55 (2010).
Google Scholar 
Liang, C., Amelung, W., Lehmann, J. & Kastner, M. Quantitative assessment of microbial necromass contribution to soil organic matter. Global Change Biol. 25, 3578–3590 (2019).ADS 

Google Scholar 
Engelking, B., Flessa, H. & Joergensen, R. G. Shifts in amino sugar and ergosterol contents after addition of sucrose and cellulose to soil. Soil Biol. Biochem. 39, 2111–2118 (2007).
Google Scholar 
Chander, K. & Joergensen, R. G. Decomposition of 14C glucose in two soils with different amounts of heavy metal contamination. Soil Biol. Biochem. 33, 1811–1816 (2001).
Google Scholar 
Zhu, Z. et al. Fate of rice shoot and root residues, rhizodeposits, and microbial assimilated carbon in paddy soil – part 2: turnover and microbial utilization. Plant Soil. 416, 243–257 (2017).
Google Scholar 
Appuhn, A. & Joergensen, R. Microbial colonisation of roots as a function of plant species. Soil Biol. Biochem. 38, 1040–1051 (2006).
Google Scholar 
Huang, Y., Liang, C., Duan, X., Chen, H. & Li, D. Variation of microbial residue contribution to soil organic carbon sequestration following land use change in a subtropical karst region. Geoderma 353, 340–346 (2019).ADS 

Google Scholar 
Liang, C. et al. Microorganisms and their residues under restored perennial grassland communities of varying diversity. Soil Biol. Biochem. 103, 192–200 (2016).
Google Scholar 
Kallenbach, C. M., Frey, S. D. & Grandy, A. S. Direct evidence for microbial-derived soil organic matter formation and its ecophysiological controls. Nat. Commun. 7, 13630 (2016).ADS 
PubMed 
PubMed Central 

Google Scholar 
Veresoglou, S. D., Chen, B. & Rillig, M. C. Arbuscular mycorrhiza and soil nitrogen cycling. Soil Biol. Biochem. 46, 53–62 (2012).
Google Scholar 
Treseder, K. K. A meta-analysis of mycorrhizal responses to nitrogen, phosphorus, and atmospheric CO2 in field studies. New Phytol. 164, 347–355 (2004).PubMed 

Google Scholar 
Xu, Y. et al. Microbial assimilation dynamics differs but total mineralization from added root and shoot residues is similar in agricultural Alfsols. Soil Biol. Biochem. 148, 107901 (2020).
Google Scholar 
Chenu, C. & Stotzky, G. Syntax of referencing in Interactions between soil particles and microorganisms (eds. Huang, P., Bollag, J. & Senesi, N.) 3–39 (Wiley-VCH, 2002).Chantigny, M., Angers, D., Prévost, D., Vézina, L.-P. & Chalifour, F. Soil aggregation and fungal and bacterial biomass under annual and perennial cropping systems. Soil Sci. Soc. Am. J. 61, 262–267 (1997).ADS 

Google Scholar 
Liang, C., Duncan, D., Balser, T., Tiedje, J. & Jackson, R. Soil microbial residue storage linked to soil legacy under biofuel cropping systems in southern Wisconsin, USA. Soil Biol. Biochem. 57, 939–942 (2013).
Google Scholar 
Feng, Y. et al. Temperature thresholds drive the global distribution of soil fungal decomposers. Glaobal Change Biol. 28, 2779–2789 (2022).
Google Scholar 
An, T. et al. Carbon fluxes from plants to soil and dynamics of microbial immobilization under plastic film mulching and fertilizer application using 13C pulse-labeling. Soil Biol. Biochem. 80, 53–61 (2015).
Google Scholar 
Lauer, F., Kösters, R., du Preez, C. C. & Amelung, W. Microbial residues as indicators of soil restoration in South African secondary pastures. Soil Biol. Biochem. 43, 787–794 (2011).
Google Scholar  More

Bolliger, J., Kienast, F. & Zimmermann, N. E. Risk of global warming on montane and subalpine forests in Switzerland—A modeling study. Reg. Environ. Change 1, 99–111 (2000).
Google Scholar 
Bugmann, H. & Pfister, Ch. Impacts of interannual climate variability on past and future forest composition. Reg. Environ. Change 1, 112–125 (2000).
Google Scholar 
Becker, A. & Bugmann, H. (eds.) Global change and mountain regions: The Mountain Research Initiative. IHDP Report 13, GTOS Report 28 and IGBP Report 49, Stockholm (2001).Kullman, L. 20th Century climate warming and tree-limit rise in the southern Scandes of Sweden. Ambio 30, 72–80. https://doi.org/10.1579/0044-7447-30.2.72 (2001).CAS 
PubMed 

Google Scholar 
Körner, Ch. & Paulsen, J. A world-wide study of high altitude treeline temperatures. J. Biogeogr. 31, 713–732. https://doi.org/10.1111/j.1365-2699.2003.01043.x (2004).
Google Scholar 
Harsch, M. A. & Bader, M. Y. Treeline form—A potential key to understanding treeline dynamics. Global Ecol. Biogeogr. 20, 582–596. https://doi.org/10.1111/j.1466-8238.2010.00622.x (2011).
Google Scholar 
Tokarczyk, N. Forest encroachment on temperate mountain meadows: scale, drivers, and current research directions. Geogr. Pol. 90, 463–480 (2017).
Google Scholar 
Vitali, A. et al. Pine recolonization dynamics in Mediterranean human-disturbed treeline ecotones. For. Ecol. Manag. 435, 28–37. https://doi.org/10.1016/j.foreco.2018.12.039 (2019).
Google Scholar 
Heikkinen, O., Obrębska-Starkel, B. & Tuhkanen, S. Introduction: the timberline—A changing battlefront. Prace Geograficzne UJ 98, 7–16 (1995).
Google Scholar 
Mattson, J. Human impact on the timberline in the far North of Europe. Zeszyty Naukowe UJ, Prace Geogr. 98, 41–56 (1995).
Google Scholar 
Stanisci, A., Lavieri, D., Acosta, A. & Blasi, C. Structure and diversity trends at Fagus timberline in central Italy. Community Ecol. 1, 133–138 (2000).
Google Scholar 
Gehrig-Fasel, J., Guisan, A. & Zimmermann, N. E. Tree line shifts in the Swiss Alps: Climate change or land abandonment?. J. Veg. Sci. 18, 571–582 (2007).
Google Scholar 
Feurdean, A. et al. Long-term land-cover/use change in a traditional farming landscape in Romania inferred from pollen data, historical maps and satellite images. Reg. Environ. Change 17, 2193–2207. https://doi.org/10.1007/s10113-016-1063-7 (2017).
Google Scholar 
Burga, C. A., Bührer, S. & Klötzli, F. Mountain ash (Sorbus aucuparia) forests of the Central and Southern Alps (Grisons and Ticino, Switzerland-Prov. Verbano-Cusio-Ossola, N-Italy): Plant ecological and phytosociological aspects. Tuexenia 39, 121–138 (2019).
Google Scholar 
Slayter, R. O. & Noble, I. R. Dynamics of Montane Treelines. In Landscape Boundaries, Consequences for Biotic Diversity and Ecological Flows. Ecological Studies Vol. 92 (eds Hansen, A. J. & di Castri, F.) 346–359 (Springer-Verlag, 1992).
Google Scholar 
Bryn, A. Recent forest limit changes in south-east Norway: Effects of climate change or regrowth after abandoned utilisation?. Nor. Geogr. Tidsskr. 62(4), 251–270. https://doi.org/10.1080/00291950802517551 (2008).
Google Scholar 
Lu, X., Liang, E., Wang, Y., Babst, F. & Camarero, J. J. Mountain treelines climb slowly despite rapid climate warming. Glob. Ecol. Biogeogr. 30(1), 305–315. https://doi.org/10.1111/geb.13214 (2021).
Google Scholar 
Armand, A. D. Sharp and Gradual Mountain Timberlines as Result of species Interaction. Landscape Boundaries, Consequences for Biotic Diversity and Ecological Flows. In Ecological Studies Vol. 92 (eds Hansen, A. J. & di Castri, F.) 360–377 (Springer-Verlag, 1992).
Google Scholar 
Kucharzyk, S. Ekologiczne znaczenie drzewostanów w strefie górnej granicy lasu w Karpatach Wschodnich i ich wrażliwość na zmiany antropogeniczne [Ecological importance of stands at the upper forest limit in the Eastern Carpathians and their sensibility to anthropogenic changes]. Roczn. Bieszcz. 14, 15–43 (2006) (in Polish with English summary).
Google Scholar 
Surina, B. & Rakaj, M. Subalpine beech forest with Hairy alpenrose (Polysticho lonchitis-Fagetum Rhododendretosum hirsuti subass. nova) on Mt. Snežnik (Liburnian Karst, Dinaric Mts). Hacquetia 6, 195–208 (2007).
Google Scholar 
Kucharzyk, S. Zmiany przebiegu górnej granicy lasu w pasmie Szerokiego Wierchu w Bieszczadzkim Parku Narodowym [Changes of upper forest limit in the Szeroki Wierch range (Bieszczady National Park)]. Roczn. Bieszcz. 12, 81–102 (2004) (in Polish with English summary).
Google Scholar 
Kucharzyk, S. & Augustyn, M. Dynamika górnej granicy lasu w Bieszczadach Zachodnich – zmiany w ciągu półtora wieku [The upper forest limit dynamics in the Western Bieszczady Mts.—Changes over a century and a half]. Stud. Nat. 54, 133–156 (2008) (in Polish with English summary).
Google Scholar 
Kubijowicz, W. Życie pasterskie w Beskidach Wschodnich [La Vie Pastorale dans les Beskides Orientales]. Prace Instytutu Geograficznego UJ 5, 3–30 (1926) (in Polish).
Google Scholar 
Zarzycki, K. Lasy Bieszczadów Zachodnich [The forests of the Western Bieszczady Mts (Polish Eastern Carpathians)]. Acta Agr. et Silv. Ser. Leśna 3, 1–131 (1963) (in Polish with English summary).
Google Scholar 
Augustyn, M. Połoniny w Bieszczadach Zachodnich [Almen im westlichen Bieszczady-Gebirge]. Materiały Muzeum Budownictwa Ludowego w Sanoku 31, 88–98 (1993) (in Polish with German summary).
Google Scholar 
Winnicki, T. Zbiorowiska roślinne połonin Bieszczadzkiego Parku Narodowego (Bieszczady Zachodnie, Karpaty Wschodnie) [Plant communities of subalpine poloninas in the Bieszczady National Park (Western Bieszczady Mts, Eastern Carpathians)]. Monogr. Bieszczadzkie 4, 1–215 (1999) (in Polish with English summary).
Google Scholar 
Mróz, W. Zróżnicowanie szaty roślinnej przy górnej granicy lasu w Bieszczadach Wschodnich i Zachodnich [The diversity of vegetation near the upper timberline in the Eastern and the Western Bieszczady Mts]. Roczn. Bieszcz. 14, 45–62 (2006) (in Polish with English summary).
Google Scholar 
Augustyn, M. & Kucharzyk, S. Górna granica lasu na terenie wsi Ustrzyki Górne i Wołosate w końcu XVIII wieku [Timberline in the Western Bieszczady Mts.]. Roczn. Bieszcz. 20, 15–27 (2012) (in Polish with English summary).
Google Scholar 
Jeník, J. Succession on the Połonina Balds in the Western Bieszczady, the Eastern Carpathians. Tuexenia 3, 207–216 (1983).
Google Scholar 
Michalik, S. & Szary, A. Zbiorowiska leśne Bieszczadzkiego Parku Narodowego [The forest communities of the Bieszczady National Park]. Monogr. Bieszcz. 1, 1–175 (1997).
Google Scholar 
Zemanek, B. & Winnicki, T. Rośliny naczyniowe Bieszczadzkiego Parku Narodowego [Vascular plants of the Bieszczady National Park]. Monogr. Bieszcz. 3, 1–249 (1999) (in Polish with English summary).
Google Scholar 
Kucharzyk, S. & Augustyn, M. Trwałość polan reglowych w Bieszczadzkim Parku Narodowym [Stability of mountain glades in the Bieszczady National Park]. Roczn. Bieszcz. 18, 45–58 (2010) (in Polish with English summary).
Google Scholar 
Durak, T., Żywiec, M. & Ortyl, B. Rozprzestrzenianie się zarośli drzewiastych w piętrze połonin Bieszczad Zachodnich [Expansion of brushwood in the subalpine zone of the Western Bieszczady Mts]. Sylwan 157, 130–138 (2013) (in Polish with English summary).
Google Scholar 
Durak, T., Żywiec, M., Kapusta, P. & Holeksa, J. Impact of land use and climate changes on expansion of woody species on subalpine meadows in the Eastern Carpathians. For. Ecol. Manag. 339, 127–135. https://doi.org/10.1016/j.foreco.2014.12.014 (2015).
Google Scholar 
Durak, T., Żywiec, M., Kapusta, P. & Holeksa, J. Rapid spread of a fleshy-fruited species in abandoned subalpine meadows—Formation of an unusual forest belt in the eastern Carpathians. iForest – Biogeosci. For. 9, 337–343. https://doi.org/10.3832/ifor1470-008 (2015).
Google Scholar 
Wężyk, P. & Hawryło, P. Analiza struktury 3D drzewostanów Bieszczadzkiego PN na podstawie danych lotniczego skanowania laserowego oraz ortofotomap lotniczych CIR [3D structure analysis of stands of the Bieszczady National Park on the basis of airborne laser scanning data and CIR aerial ortho-photomaps] (ProGea Consulting, 2015) (in Polish).Anselin, L. Local indicators of spatial association—LISA. Geogr. Anal. 27, 93–115. https://doi.org/10.1111/j.1538-4632.1995.tb00338.x (1995).
Google Scholar 
Scott, L. M. & Janikas, M. V. Spatial Statistics in ArcGIS. In Handbook of Applied Spatial Analysis (eds Fischer, M. M. & Getis, A.) 27–41 (Springer, 2010).
Google Scholar 
Cui, H., Wu, L., Hu, S., Lu, R. & Wang, S. Research on the driving forces of urban hot spots based on exploratory analysis and binary logistic regression model. Trans. GIS 25(3), 1522–1541. https://doi.org/10.1111/tgis.12739 (2021).
Google Scholar 
Pierce, K. B., Lookingbill, T. & Urban, D. A simple method for estimating potential relative radiation (PRR) for landscape-scale vegetation analysis. Landsc. Ecol. 20, 137–147 (2005).
Google Scholar 
Riley, S. J., DeGloria, S. D. & Elliot, R. A terrain ruggedness index that quantifies topographic heterogeneity. Int. J. Sc. 5, 23–27 (1999).
Google Scholar 
Böhner, J. & Antonić, O. Land-surface parameters specific to topo-climatology. Geomorphometry – Concepts, Softw. Appl. Dev. Soil Sci. 33, 195–226. https://doi.org/10.1016/S0166-2481(08)00008-1 (2009).
Google Scholar 
R Core Team. R: A Language and Environment for Statistical Computing (R Core Team, 2021).
Google Scholar 
Agresti, A. An Introduction to Categorical Data Analysis 2nd edn. (Wiley & Sons Inc., 2007).MATH 

Google Scholar 
Cottrell, A. Gnu Regression, Econometrics and Time-series Library gretl. http://gretl.sourceforge.net/(2020).Hellevik, O. Linear versus logistic regression when the dependent variable is a dichotomy. Qual. Quant. 43, 59–74 (2009).
Google Scholar 
Azen, R. & Traxel, N. Using dominance analysis to determine predictor importance in logistic regression. J. Educ. Behav. Stat. 34, 319–347. https://doi.org/10.3102/1076998609332754 (2009).
Google Scholar 
Borcard, P., Legendre, P. & Drapeau, P. Partialling out the spatial component of ecological variation. Ecology 73, 1045–1055 (1992).
Google Scholar 
Przybylska, K. & Kucharzyk, S. Skład gatunkowy i struktura lasów Bieszczadzkiego Parku Narodowego [Species composition and structure of forest of the Bieszczady National Park. Monogr. Bieszcz. 6, 1–159 (1999) (in Polish with English summary).
Google Scholar 
Bader, M. Y. et al. A global framework for linking alpine-treeline ecotone patterns to underlying processes. Ecography 44(2), 265–292. https://doi.org/10.1111/ecog.05285 (2021).
Google Scholar 
Nowosad, M. Zarys klimatu Bieszczadzkiego Parku Narodowego i jego otuliny w świetle dotychczasowych badań [Outlines of climate of the Bieszczady National Park and its bufferzone in the light of previous studies]. Roczn. Bieszcz. 4, 163–183 (1995) (in Polish with English summary).
Google Scholar 
Nowosad, M. & Wereski, S. Warunki klimatyczne. Bieszczadzki Park Narodowy–40 lat ochrony [Climatic conditions. Bieszczady National Park–40 years of protection]. In Bieszczadzki Park Narodowy [The Bieszczady National Park] (eds Górecki, A. & Zemanek, B.) 31–38 (Wyd. Bieszczadzki Park Narodowy, 2016) (in Polish with English summary).
Google Scholar 
Kukulak, J. Neotectonics and planation surfaces in the High Bieszczady Mountains (Outer Carpathians, Poland). Ann. Soc. Geol. Pol. 74, 339–350 (2004).
Google Scholar 
Haczewski, G., Kukulak, J. & Bąk, K. Budowa geologiczna i rzeźba Bieszczadzkiego Parku Narodowego [Geology and relief of the Bieszczady National Park]. Prace monograficzne (Akademia Pedagogiczna im. Komisji Edukacji Narodowej w Krakowie) 468, 1–156 (2007) (in Polish with English summary).
Google Scholar 
Skiba, S., Drewnik, M., Kacprzak, A. & Kołodziejczyk, M. Gleby litogeniczne Bieszczadów i Beskidu Niskiego [Lithogenous soils of the Bieszczady and Beskid Niski Mts (Polish Carpathians)]. Roczn. Bieszcz. 7, 387–396 (1998) (in Polish with English summary).
Google Scholar 
Skiba, S. & Winnicki, T. Gleby zbiorowisk roślinnych bieszczadzkich połonin [Soils of the subalpine meadows plant communities in the Bieszczady Mts]. Roczn. Bieszcz. 4, 97–109 (1995) (in Polish with English summary).
Google Scholar 
Musielok, Ł, Drewnik, M., Szymański, W. & Stolarczyk, M. Classification of mountain soils in a subalpine zone—A case study from the Bieszczady Mountains (SE Poland). Soil Sci. Annu. 70, 170–177. https://doi.org/10.2478/ssa-2019-0015 (2019).CAS 

Google Scholar 
Spatz, G. Succession patterns on mountain pastures. Vegetatio 43, 39–41 (1980).
Google Scholar 
Kozak, J. Zmiany powierzchni lasów w Karpatach Polskich na tle innych gór świata [Changes in the Land Cover in the Polish Carpathians at the Turn of the 20th and 21st Century in Relation to Local Development Level]. Wydawnictwo Uniwersytetu Jagiellońskiego, Kraków (2005) (in Polish with English summary).Vitali, A., Urbinati, C., Weisberg, P. J., Urza, A. K. & Garbarino, M. Effects of natural and anthropogenic drivers on land-cover change and treeline dynamics in the Apennines (Italy). J. Veg. Sci. 29(2), 189–199. https://doi.org/10.1111/jvs.12598 (2018).
Google Scholar 
Micu, D. M., Dumitrescu, A., Cheval, S., Nita, I.-A. & Birsan, M.-V. Temperature changes and elevation-warming relationships in the Carpathian Mountains. Int. J. Climatol. 41, 2154–2172. https://doi.org/10.1002/joc.6952 (2020).
Google Scholar 
Rehman, A. Ziemie dawnej Polski. Cz. I. Karpaty [The lands of ancient Poland. Part I. The Carpathians]. (Gubrynowicz i Schmidt, Lwów) (1895) (in Polish).Frey, W. The influence of snow on growth and survival of planted trees. Arct. Alp. Res. 15, 241–251 (1983).
Google Scholar 
Malanson, G. P. et al. Alpine treeline of Western North America: Linking organism-to-landscape dynamics. Phys. Geogr. 28, 378–396. https://doi.org/10.2747/0272-3646.28.5.378 (2007).
Google Scholar 
Holtmeier, F. K. & Broll, G. Wind as an ecological agent at treelines in North America, the Alps, and the European Subarctic. Phys. Geogr. 31, 203–233. https://doi.org/10.2747/0272-3646.31.3.203 (2010).
Google Scholar 
Barclay, A. M. & Crawford, R. M. M. Winter desiccation stress and resting bud viability in relation to high altitude survival in Sorbus aucuparia L. Flora 172, 21–34 (1982).
Google Scholar 
Raspé, O., Findlay, C. & Jacquemart, A. L. Sorbus aucuparia L. J. Ecol. 88, 910–930 (2000).
Google Scholar 
Zerbe, S. On the ecology of Sorbus aucuparia (Rosaceae) with special regard to germination, establishment and growth. Pol. Bot. J. 46, 229–239 (2001).
Google Scholar 
Smith, W. K., Germino, M. J., Hancock, T. E. & Johnson, D. M. Another perspective on altitudinal limits of alpine timberlines. Tree Physiol. 23, 1101–1112 (2003).PubMed 

Google Scholar 
Trant, A., Higgs, E. & Starzomski, B. M. A century of high elevation ecosystem change in the Canadian Rocky Mountains. Sci. Rep. 10, 9698. https://doi.org/10.1038/s41598-020-66277-2 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Barbeito, I., Dawes, M. A., Rixen, C., Senn, J. & Bebi, P. Factors driving mortality and growth at treeline: A 30-year experiment of 92 000 conifers. Ecology 93(2), 389–401 (2012).PubMed 

Google Scholar 
Kullman, L. A 25-year survey of geoecological change in the scandes mountains of Sweden. Geogr. Ann. Ser. B 79, 139–165 (1997).
Google Scholar 
Pękala, K. Rzeźba Bieszczadzkiego Parku Narodowego [Relief of the Bieszczady National Park]. Roczn. Bieszcz. 6, 19–38 (1997) (in Polish with English summary).
Google Scholar 
Kullman, L. Temporal and spatial aspects of subalpine populations of Sorbus aucuparia in Sweden. Ann. Bot. Fenn. 23, 267–275 (1986).
Google Scholar 
Hoersch, B. Modelling the spatial distribution of montane and subalpine forests in the Central Alps using digital elevation models. Ecol. Model. 168, 267–282 (2003).
Google Scholar 
Resler, L. M., Butler, D. R. & Malanson, G. P. Topographic shelter and conifer establishment and mortality in an alpine environment, Glacier National Park, Montana. Phys. Geogr. 26, 112–125 (2005).
Google Scholar 
Kollmann, J. Regeneration window for fleshy-fruited plants during scrub development on abandoned grassland. Ecoscience 2, 213–222 (1995).
Google Scholar 
Lediuk, K. D., Damascos, M. A., Puntieri, J. G. & de Torres Curth, M. I. Population dynamics of an invasive tree, Sorbus aucuparia, in the understory of a Patagonian forest. Plant Ecol. 217, 899–911 (2016).
Google Scholar 
McCutchan, M. H. & Fox, D. G. Effect of elevation and aspect on wind, temperature and humidity. J. Appl. Meteorol. Climatol. 25(12), 1996–2013 (1986).ADS 

Google Scholar 
Stage, A. R. & Salas, C. Interactions of elevation, aspect, and slope in models of forest species composition and productivity. For. Sci. 53, 486–492 (2007).
Google Scholar 
Pocewicz, A. L., Gessler, P. & Robinson, A. P. The relationship between effective plant area index and Landsat spectral response across elevation, solar insolation, and spatial scales in a northern Idaho forest. Can. J. For. Res. 34, 465–480 (2004).
Google Scholar 
Kucharzyk, S. & Sugiero, D. Zróżnicowanie dynamiki procesów lasotwórczych w buczynach bieszczadzkich w zależności od wystawy i wzniesienia [Variability of the dynamics of forest development processes in the Bieszczady beech forests in relation to exposition and altitude]. Sylwan 7, 29–38 (2007) (in Polish with English summary).
Google Scholar 
Drewnik, M., Musielok, Ł, Stolarczyk, M., Mitka, J. & Gus, M. Effects of exposure and vegetation type on organic matter stock in the soils of subalpine meadows in the Eastern Carpathians. CATENA 147, 167–176. https://doi.org/10.1016/j.catena.2016.07.014 (2016).CAS 

Google Scholar 
Zheng, L. et al. Tree regeneration patterns on contrasting slopes at treeline ecotones in Eastern Tibet. Forests 12, 1605. https://doi.org/10.3390/f12111605 (2021).
Google Scholar  More

The studied viviparous clausiliids developed four types of morphological adaptations that facilitate the delivery of embryos through the shell aperture: (1) reduction of the clausiliar apparatus, (2) decrease of embryonic shell width, (3) widening of the shell canal, and (4) development of a flexible embryonic shell.Reduction of the clausiliar apparatusMembers of the Reinia genus, arboreal species from Japan (Fig. 1), show the most advanced adaptations to live-bearing compared to hypothetical ancestral Phaedusinae. The shell shape in these species is more conical than fusiform, the number of whorls decreases, and the aperture widens. One of the species, R. variegata, features almost full reduction of the clausiliar apparatus that consists of only vestigial folds (Fig. 1F). This species also lacks the clausilium, so the entrance through the aperture is unprotected.Figure 1Different stages of reduction of apertural barriers in members of genus Reinia: R. ashizuriensis (A–C; upper row) and R. variegata (D–F; lower row). (A,D) Adult shells; (B,C,E,F) adult shells with body whorl cut open dorsally in microCT visualisation. cp clausilium plate, il inferior lamella, pr principal plica, sc subcolumellar lamella, sl superior lamella, sp spiral lamella, upp upper palatal plica.Full size imageDecrease of embryonic shell widthAnother adaptation concerns the shape of the embryonic shell (“protoconch”), which becomes very narrow in some viviparous species. This feature is conspicuous because embryonic whorls remain in the adult shell as apical whorls. For instance in S. addisoni (Fig. 2A–D), the apical part being much narrower than the first whorls of the teleoconch is a clear evidence that the growth trajectory has changed abruptly after birth. Other examples include E. cylindrella and E. steetzneri, in which both the protoconch and the teleoconch are very narrow, yet at the borderline between these parts, the shell axis is slightly bent (Fig. 2E–L). We suppose that this feature develops as a result of obstruction during birth.Figure 2Width difference between protoconch and teleoconch in Stereophaedusa addisoni (A–D, upper row), Euphaedusa cylindrella (E–H, middle row), Euphaedusa steetzneri (I–L, lower row). (A,C,E,G,I,K) Adult shells with very narrow apical whorls; (B,F,J) X-rayed adults; (F,J) with retained embryos inside; (D,H,L) X-rays of apical part of adult shell with schematic drawings of a neonate.Full size imageWidening of the shell canalThe third type of adaptation is the widening of the shell canal in the body whorl, allowing for easier passage of the embryo between the lamellae and plicae of the apertural barriers. In this case, the outline of the shell changes only slightly giving the body whorl a more convex appearance. A substantial difference to egg-laying species concerns the apertural barriers: the clausiliar includes a broad clausilium plate and a spirally ascending inferior lamella (Fig. 3A–D). These modifications result in a spacious shell canal in the body whorl, for example in S. addisoni and E. sheridani, that can accommodate the transfer of a large embryo. Table 1 presents neonatal size in these species (shell width ca. 1.2 mm), which is very similar to their clausilium width (ca. 1.1–1.2 mm).Figure 3Two types of clausiliar apparatus occurring in Phaedusinae in microCT visualisation: with spirally ascending inferior lamella and wide clausilium plate (upper row), and with straight ascending inferior lamella and narrow clausilium plate (lower row). (A) T. sheridani adult shell with the body whorl cut open dorsally; (B) clausilium of T. sheridani; (C) clausilium of S. addisoni; (D) clausilium of R. ashizuriensis; (E) Zaptyx ventriosa adult shell with body whorl cut open dorsally; (F) clausilium of Z. ventriosa; (G,H) clausilia of O. miranda. Note, that all depicted species are viviparous.Full size imageTable 1 Shell size of studied Phaedusinae species.Full size tableMost viviparid clausiliids develop one of these three types of modification; some adaptations co-occur within a single species, for example a wide clausilium accompanies a narrow apex. Interestingly, the Reinia genus includes taxa with a gradual escalation of viviparity-related adaptations: R. ashizurensis, with a stout shell shape and a low number of whorls, has fully developed apertural barriers with a broad clausilium plate (Fig. 1A–C), while its congener, R. variegata, has reduced apertural barriers (Fig. 1D–F).Development of a flexible embryonic shellThe fourth type of adaptation found in Phaedusinae concerns the structure of the embryonic shells. We report this adaptation in O. miranda and Z. ventriosa.Oospira miranda is a dextral, often decollated, ground-dwelling species from Vietnam (Fig. 4A). The species is viviparous: during microCT scanning of museum specimens, we found embryos within a parental shell (Fig. 4B); in laboratory culture, we observed neonates immediately after live birth (Fig. 4C,D). Morphological characters recognized in the adult shell, i.e., a wide apex (= wide embryonic shell), straightly ascending inferior lamella, and a narrow clausilium plate (Fig. 3G,H), seemed to exclude the possibility of live-bearing reproduction, as embryos are too large to pass through the shell canal at the narrowest point. The height and width of the neonatal shell (mean values: 5.19 mm, 3.59 mm) evidently exceeds the width of the clausilium plate in this species (1.97 mm) (Table 1). However, under closer examination, we found the shell to be thin and delicate, which we refer to as a ‘soft shell’. In direct examination, the neonatal shell of O. miranda resembles cellophane, which may keep a given shape for a long time but becomes distorted already under slight pressure.Figure 4Viviparous clausiliids and their ‘soft-shelled’ neonates born in laboratory culture. (A–D) O. miranda: adult shell, X-rayed shell with embryo visible inside, neonates; (E–H) Z. ventriosa: adult shell, X-rayed shell with eggs visible inside, neonates.Full size imageA similar adaptation exists in Z. ventriosa, a Taiwanese species with a very wide apex, never decollated, a straight ascending inferior lamella, and a narrow clausilium plate (Figs. 3E,F, 4E,F). This species produces neonates in laboratory culture (Fig. 4G–H). The dimensions of the neonates (mean values: height 3.37 mm, width 2.51 mm) exceed at last twofold the width of the clausilium plate (1.08 mm). The shells of such freshly delivered juveniles, when gently touched with laboratory tweezers, became dented, but not fractured. More intense and stronger pressing can break this dentation.These initial observations, that we made during the maintenance of the laboratory culture, suggested that the neonatal shells of O. miranda and Z. ventriosa have flexible walls. These ‘soft-shells’ seem to be highly malleable during the entire embryonic development period and delivery through apertural barriers, hardening shortly after birth. We further investigated the physical properties of the embryonic shell by means of microcomputed tomography and scanning electron microscopy.Microcomputed tomographyWe scanned ‘soft-shelled’ neonates of O. miranda and Z. ventriosa, together with ‘hard-shelled’ embryos and neonates of S. addisoni and T. sheridani, in order to compare the density and thickness of the shells (Fig. 5).Figure 5Comparison of embryonic shell thickness in clausiliids: ‘soft-shelled’ neonates of Z. ventriosa (A,B,G,H) and O. miranda (C,D,I,J); “hard-shelled” neonate of S. addisoni (E,K) and embryo of T. sheridani (F,L) scanned inside a parental shell. Upper row—microCT visualisation of shell surface; middle row—microCT sections of those specimens; (M–O) X-ray photographs of S. addisoni (embryo from dissected adult) and Z. ventriosa (neonate) enlarged in (N,O), respectively, showing the difference in shell density and thickness; (P) microCT based volume rendering of O. miranda (left) and S. addisoni (right) neonates, showing difference between relative density of their shells.Full size imagePreliminary observations using the two-dimensional X-ray photographs showed a difference in thickness and density between S. addisoni and Z. ventriosa (Fig. 5M, enlarged in N and O, respectively). The 3D visualization of O. miranda and S. addisoni (the same microCT scanning and reconstruction parameters) confirmed the difference between density and shell thickness of these two species (Fig. 5P).Due to variations in wall thickness within the neonatal shell (e.g., between the first and the second whorls), it is not possible to precisely determine the thickness of the shell wall. The accuracy of the measurement is also limited by the resolution of the microCT scans, especially in the case of the relatively large neonates of O. miranda and Z. ventriosa. When scanning the whole embryonic shell of Z. ventriosa (approximately 3.5 mm in height), the size of the voxel was approximately 1 µm. Thus, we cannot determine the shell thickness down to the nearest micron, but we can estimate it from a few to a dozen microns. A direct comparison between virtual microCT sections of specimens scanned under the same conditions shows a clear difference between the ‘soft-shelled’ and ‘hard-shelled’ taxa (Fig. 5G–L). The ’hard-shelled’ neonates have a shell wall of 30–40 µm thick. We examined the sequence of three ’soft-shelled’ O. miranda specimens that differed in size (the exact time of birth of each of the cultured neonates is unknown, ca. 1–2 days). The larger (older) the neonate was, the thicker the shell. The shell of the largest of the studied O. miranda was up to 20 µm thick. However, the shell wall of this relatively large juvenile (several millimeters in height) still did not reach the thickness of the small ’hard-shelled’ T. sheridani embryo, which was already about 30–40 µm thick, stiff and rigid during the retention in the genital tract. The neonates of O. miranda and Z. ventriosa were much larger than the embryos and neonates of S. addisoni and R. variegata (Table 1), however, the former taxa has much thinner shells.Scanning electron microscopyAfter the non-invasive microCT scan, we scanned embryos and neonates using SEM (Fig. 6). The different properties of the shells of Z. ventriosa and O. miranda vs. S. addisoni and R. variegata were already visible during the preparation of the analysis. Under vacuum conditions, the soft shells of Z. ventriosa and O. miranda shrank and crumpled, creating a cellophane-like surface (Fig. 6A). Embryos and neonates of S. addisoni and R. variegata did not require any special preparation and their shell shape remained unchanged under the vacuum conditions applied during the SEM examination (Fig. 6D,E). To reduce the shell deformations, we freeze-dried the next group of thin-shelled neonates prior to SEM analyses (Fig. 6B,C).Figure 6Neonates of O. miranda (A,B,F,I,L,M,O) and Z. ventriosa (C,G,J,P) in direct comparison with hard-shelled embryos and neonates of R. variegata (D,N,Q) and S. addisoni (E,H,K); SEM microphotographs. The vacuum conditions in SEM led to the shrinkage of the thin O. miranda shell (A); freeze-drying of ‘soft-shelled’ neonates prior to SEM imaging reduced the level of deformity (B,C). Contrastingly, R. variegata and S. addisoni shells do not require special preparation and retain their shape (D,E). (F) The dented surface of O. miranda neonate and SEM-close-up (I) on a cross-section of the shell just a few micrometers thick (arrow in F indicates the region enlarged in I). (G,J) Shell of Z. ventriosa in comparison with similarly ornamented fragment of S. addisoni (H,K); note several times thicker shell in the latter (arrows in G,H indicate the regions enlarged in J,K, respectively). (L,M) Inner surface of intact periostracum which still connects two fragments of broken aragonite shell of O. miranda (the arrow in M indicates the region enlarged in L); note the difference between shell thickness in O. miranda (L,M) and R. variegata (N). All observed specimens have similar crossed-lamellar microstructure (L–Q). However, just as shell thickness, also the number of lamellar layers of alternate orientation within the shell differs (L,M,O,P vs N,Q).Full size imageThe SEM studies allowed for complementary measurements of the shells. In the broken fragments of Z. ventriosa and O. miranda, the thickness of the shell wall ranged from 2–3 µm (Fig. 6F,G,I,J,L,M) to 18 µm in the largest neonate of O. miranda (Fig. 6O). The shells of S. addisoni (Fig. 6H,K) and R. variegata (Fig. 6N) are several times thicker.All analyzed samples have a thin ( More

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A roaring trans-European herring trade that began in the Viking Age might have depleted stocks1.

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doi: https://doi.org/10.1038/d41586-022-03431-y

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Roff, D. A. The Evolution of Life Histories: Theory and Analysis (Chapman and Hall, 1992).
Google Scholar 
Sæther, B. E. & Bakke, O. Avian life history variation and contribution of demographic traits to the population growth rate. Ecology 81, 642–653 (2000).
Google Scholar 
Koons, D. N., Pavard, S., Baudisch, A. & Metcalf, J. E. C. Is life-history buffering or lability adaptive in stochastic environments?. Oikos 118, 972–980 (2009).
Google Scholar 
Boyce, M. S., Haridas, C. V. & Lee, C. T. Demography in an increasingly variable world. Trends Ecol. Evol. 21, 141–148 (2006).PubMed 

Google Scholar 
Morris, W. F. & Doak, D. F. Buffering of life histories against environmental stochasticity: Accounting for a spurious correlation between the variabilities of vital rates and their contributions to fitness. Am. Nat. 163, 579–590 (2004).PubMed 

Google Scholar 
Ripple, W. J. et al. Saving the world’s terrestrial megafauna. Bioscience 66(10), 807–812 (2016).PubMed 
PubMed Central 

Google Scholar 
He, F. et al. Disappearing giants: A review of threats to freshwater megafauna. WIREs Water 4, e1208 (2017).
Google Scholar 
Blackburn, T. M., Cassey, P., Duncan, R. P., Evans, K. L. & Gaston, K. J. Avian extinction and mammalian introductions on oceanic islands. Science 305(5692), 1955–1958 (2004).ADS 
CAS 
PubMed 

Google Scholar 
Courchamp, F., Langlais, M. & Sugihara, G. Rabbits killing birds: Modelling the hyperpredation process. J. Anim. Ecol. 69, 154–164 (2000).
Google Scholar 
Roemer, G. W., Coonan, T. J., Garcelon, D. K., Bascompte, J. & Laughrin, L. Feral pigs facilitate hyperpredation by golden eagles and indirectly cause the decline of the island fox. Anim. Conserv. 4, 307–318 (2001).
Google Scholar 
Kristan, W. B. & Boarman, W. I. Spatial patterns of risk of common raven predation on desert tortoises. Ecology 84, 2432–2443 (2003).
Google Scholar 
Whelan, C. J., Brown, J. S. & Maina, G. Search biases, frequency-dependent predation and species co-existence. Evol. Ecol. Res. 5, 329–343 (2003).
Google Scholar 
Moleón, M., Almaraz, P. & Sánchez-Zapata, J. A. An emerging infectious disease triggering large-scale hyperpredation. PLoS ONE 3, e2307 (2008).ADS 
PubMed 
PubMed Central 

Google Scholar 
Moleón, M., Almaraz, P. & Sánchez-Zapata, J. A. Inferring ecological mechanisms from hunting bag data in wildlife management: A reply to blanco-aguiar et al. 2012. Eur. J. Wildl. Res. 59, 599–608 (2013).
Google Scholar 
Bate, A. M. & Hilker, F. M. Rabbits protecting birds: Hypopredation and limitations of hyperpredation. J. Theor. Biol. 297, 103–115 (2012).ADS 
MathSciNet 
PubMed 
MATH 

Google Scholar 
Turner, F. B., Medica, P. A. & Lyons, C. L. Reproduction and survival of the desert tortoise (Scaptochelys agassizii) in Ivanpah Valley California. Copeia 1984(4), 811–820 (1984).
Google Scholar 
Graciá, E. et al. Assessment of the key evolutionary traits that prevent extinctions in human altered habitats using a spatially explicit individual-based model. Ecol. Model. 415, 108823 (2020).
Google Scholar 
Segura, A., Jiménez, J. & Acevedo, P. Predation of young tortoises by rabbits: The effect of habitat structure on tortoise detectability and abundance. Sci. Rep. 10, 1–9 (2020).
Google Scholar 
Watson, J. The golden eagle (Bloomsbury Publishing, 2010).
Google Scholar 
Fischer, W., Zenker, D. & Baumgart, W. Ein beitrag zum bestand und zur ernährung des steinadlers Aquila chrysaetos af der balkanhalbinsel. Beiträge zur Vogelskunde 21, 275–287 (1975).
Google Scholar 
Delibes, M., Calderón, J. & Hiraldo, F. Selección de presa y alimentación en españa del águila real (Aquila chrysaetos). Ardeola 21, 285–303 (1975).
Google Scholar 
Handrinos, G. The Golden Eagle in Greece. Actes 1er Coll. Intern. Aigle Royal en Europe, Arvieux, 1986: 18–22 (1987).Bautista, J., Gil-Sánchez, J. M. & Moleón, M. Dieta del águila real en el sur de españa. Quercus 364, 17–23 (2016).
Google Scholar 
Bautista, J., Castillo, S., Paz, J. L., Llamas, J. & Ellis, D. H. Golden eagles (Aquila chrysaetos) as potential predators of barbary macaques (Macaca sylvanus) in northern Morocco: Evidences of predation. Go-South Bull. 15, 172–179 (2018).
Google Scholar 
Kouzmanov, G., Stoyanov, R. & Todorov, V. Sur la biologie et la Protection de l`Aigle royal Aquila chrysaetos en Bulgarie. In Eagle studies (eds Meyburg, B. & Chancellor, R.) 505–516 (World Working Group on Birds of Prey, 1996).
Google Scholar 
Capper, S. The predation of Testudo spp. By Golden Eagles Aquila chrysaetos in Dadia Forest Reserve, NE Greece. University of Reading (1998).Karyakin, I. V., Kovalenko, A. V., Levin, A. S. & Pazhenkov, A. S. Eagles of the Aral-Caspian region Kazakhstan. Raptors Conserv. 22, 92–152 (2011).
Google Scholar 
Papageorgiou, N., Vlachos, C., Bakaloudis, D. E., Kazaklis, A., Birtsas, P. Study on the biology and management of raptors in Dadia forest–Evros. Thessaloniki, GR (1995).Sidiropoulos, L. et al. Pronounced seasonal diet diversity expansion of golden eagles (Aquila chrysaetos) in Northern Greece during the non-breeding season: The role of tortoises. Diversity 14(2), 135 (2022).
Google Scholar 
IUCN. The IUCN red list of threatened species. Version 2020–3 (2020).Graciá, E. et al. Expansion after expansion: dissecting the phylogeography of the widely distributed spur-thighed tortoise, Testudo graeca (Testudines: Testudinidae). Biol. J. Linn. Soc. 121, 641–654 (2017).
Google Scholar 
Graciá, E. et al. Genetic patterns of a range expansion: The spur-thighed tortoise Testudo graeca graeca in southeastern Spain. Amphib. Reptil. 32, 49–61 (2011).
Google Scholar 
Graciá, E. et al. The uncertainty of late pleistocene range expansions in the western Mediterranean: A case study of the colonization of south-eastern Spain by the spur-thighed tortoise, Testudo graeca.. J. Biogeogr 40, 323–334 (2013).
Google Scholar 
Anadón, J. D., Giménez, A., Perez, I., Martinez, M. & Esteve-Selma, M. A. Habitat selection by the spur-thighed tortoise Testudo graeca in a multisuccessional landscape: implications for habitat management. Biodivers. Conserv. 15, 2287–2299 (2006).
Google Scholar 
Rodríguez-Caro, R. C. et al. Low tortoise abundances in pine forest plantations in forest-shrubland transition areas. PLoS ONE 12, e0173485 (2017).PubMed 
PubMed Central 

Google Scholar 
Rodríguez-Caro, R. C. et al. The limits of demographic buffering in coping with environmental variation. Oikos 130(8), 1346–1358 (2021).
Google Scholar 
Rodríguez-Caro, R. C., Lima, M., Anadón, J. D., Graciá, E. & Giménez, A. Density dependence, climate and fires determine population fluctuations of the spur-thighed tortoise, Testudo graeca. J. Zool. 300, 265–273 (2016).
Google Scholar 
Rodríguez-Caro, R. C. et al. A low cost approach to estimate demographic rates using inverse modeling. Biol. Conserv. 237, 358–365 (2019).
Google Scholar 
Jiménez-Franco, M. V. et al. Sperm storage reduces the strength of the mate-finding allee effect. Ecol. Evol. 10(4), 1938–1948 (2020).PubMed 
PubMed Central 

Google Scholar 
Graciá, E. et al. From troubles to solutions: Conservation of mediterranean tortoises under global change. Basic Appl. Herpetol. 34, 5–16 (2020).
Google Scholar 
Pérez, I. et al. Exurban sprawl increases the extinction probability of a threatened tortoise due to pet collections. Ecol. Model. 245, 19–30 (2012).
Google Scholar 
Del Moral, J. C. El águila real en España. Población reproductora en 2008 y método de censo. SEO/BirdLife. Madrid. pp. 30–50 (2009).Virgós, E., Cabezas-Díaz, S. & Lozano, J. Is the wild rabbit (Oryctolagus cuniculus) a threatened species in Spain? Sociological constraints in the conservation of species. Biodivers. Conserv. 16, 3489–3504 (2007).
Google Scholar 
Fernández, C. Effect of the viral haemorrhagic pneumonia of the wild rabbit on the diet and breeding success of the golden eagle Aquila chrysaetos (L.). Rev. Ecol. Terre et Vie 48, 323–329 (1993).
Google Scholar 
Villafuerte, R., Luco, D. F., Gortázar, C. & Blanco, J. C. Effect on red fox litter size and diet after rabbit haemorrhagic disease in northeastern Spain. J. Zool. 240, 764–767 (1996).
Google Scholar 
Martínez, J. A. & Zuberogoitia, I. The response of eagle owl (Bubo bubo) to an outbreak of the rabbit haemorrhagic disease. J. Ornithol. 142, 204–211 (2001).
Google Scholar 
Moleón, M. et al. Large-scale spatiotemporal shifts in the diet of a predator mediated by an emerging infectious disease of its main prey. J. Biogeogr. 36, 1502–1515 (2009).
Google Scholar 
Adamakopoulos, T., Gatzoyannis, S., Poirazidis, K. Study on the assessment, the enhancement of the legal infrastructure and the management of the protected area in the forest of Dadia. Specific Environmental Study, WWF-Greece, Athens (1995).Delibes, M., Hiraldo, F. The rabbit as prey in the Iberian Mediterranean ecosystem. In Proceedings of the World Lagomorph Conference. Guelph: University of Guelph. 1979: 614–622 (1979).Futuyma, D. J. & Moreno, G. The evolution of ecological specialization. Annu. Rev. Ecol. Syst. 19, 207–233 (1988).
Google Scholar 
Moleón, M. et al. Predator–prey relationships in a mediterranean vertebrate system: Bonelli’s eagles, rabbits and partridges. Oecologia 168, 679–689 (2012).ADS 
PubMed 

Google Scholar 
Fedriani, J. M., Ferreras, P. & Delibes, M. Dietary response of the Eurasian badger, Meles meles, to a decline of its main prey in the Doñana national park. J. Zool. 245, 214–218 (1998).
Google Scholar 
Ferrer, M. & Negro, J. J. The near extinction of two large European predators: Super specialists pay a price. Conserv. Biol. 18, 344–349 (2004).
Google Scholar 
Lozano, J., Moleón, M. & Virgós, E. Biogeographical patterns in the diet of the wildcat, Felis silvestris Schreber, in Eurasia: Factors affecting the trophic diversity. J. Biogeogr. 33, 1076–1085 (2006).
Google Scholar 
Burgos, T. et al. Prey density determines the faecal-marking behaviour of a solitary predator, the Iberian lynx (Lynx pardinus). Ethol. Ecol. Evol. 31, 219–230 (2019).
Google Scholar 
Ontiveros, D. & Pleguezuelos, J. M. Influence of prey densities in the distribution and breeding success of Bonelli’s eagle (Hieraaetus fasciatus): Management implications. Biol. Conserv. 93, 19–25 (2000).
Google Scholar 
Araújo, M. S. & Gonzaga, M. O. Individual specialization in the hunting wasp Trypoxylon (Trypargilum) albonigrum (Hymenoptera, Crabronidae). Behav. Ecol. Sociobiol. 61, 1855–1863 (2007).
Google Scholar 
Stephens, D. W. & Krebs, J. R. Foraging Theory 1st edn. (Monographs in Behavior and Ecology. Princeton University Press, 1986).
Google Scholar 
Heath, J. A. et al. Golden Eagle dietary shifts following wildfire and shrub loss have negative consequences for nestling survivorship. Ornithol. Appl. 123(4), duabo34 (2021).
Google Scholar 
Anadón, J. D., Wiegand, T. & Giménez, A. Individual-based movement models reveal sex-biased effects of landscape fragmentation on animal movement. Ecosphere 3, 1–32 (2012).
Google Scholar 
Sanz-Aguilar, A., Anadón, J. D., Giménez, A., Ballestar, R. & Oro, D. Coexisting with fire: The case of the terrestrial tortoise Testudo graeca in mediterranean shrublands. Biol. Conserv. 144, 1040–1049 (2011).
Google Scholar 
Arroyo, B. Águila real – Aquila chrysaetos. In: Enciclopedia Virtual de los Vertebrados Españoles. Salvador, A., Morales, M. B. (Eds.). Museo Nacional de Ciencias Naturales, Madrid. http://www.vertebradosibericos.org/ (2017).Arroyo, B., Ferreiro, E., Garza, V. El águila real (Aquila chrysaetos) en España. Censo, distribución, reproducción y conservación. Serie Técnica, ICONA. Madrid (1990).Bautista, J., Gil-Sánchez, J. M., González Miras, E., Gómez, G. J. & Sánchez Balsera, J. L. Increase in the population of golden eagle in andalusian baetic system mountain ranges (southern of Spain): evidences of competition with the Bonelli’s eagle. Quercus 332, 16–22 (2013).
Google Scholar 
Rodríguez-Caro, R. C., Graciá, E., Anadón, J. D. & Giménez, A. Maintained effects of fire on individual growth and survival rates in a spur-thighed tortoise population. Eur. J. Wildl. Res. 59, 911–913 (2013).
Google Scholar 
Beissinger, S. R. & McCullough, D. R. Population viability analysis (University of Chicago Press, 2002).
Google Scholar 
Tylianakis, J. M., Didham, R. K., Bascompte, J. & Wardle, D. A. Global change and species interactions in terrestrial ecosystems. Ecol. Lett. 11, 1351–1363 (2008).PubMed 

Google Scholar 
Real, J. Biases in diet study methods in the Bonelli’s eagle. J. Wildl. Manag. 60(3), 632–638 (1996).
Google Scholar 
Moleón, M. et al. Laying the foundations for a human-predator conflict solution: Assessing the impact of Bonelli’s eagle on rabbits and partridges. PLoS ONE 6, e22851 (2011).ADS 
PubMed 
PubMed Central 

Google Scholar 
Esteve-Selma, M. A., et al. Effects of climate change on the potential distribution of Testudo graeca in southeastern Iberian Peninsula. In Graciá E, Rodríguez-Caro RC and Giménez A. Conservation of Mediterranean tortoises under global change. Madrid. Asociación Herpetológica Española. ISBN: 978-84-921999-6-9.Anadón, J. D., Giménez, A., Ballestar, R. & Pérez, I. Evaluation of local ecological knowledge as a method for collecting extensive data on animal abundance. Conserv. Biol. 23, 617–625 (2009).PubMed 

Google Scholar 
Abad, V. Variaciones del Índice corporal en una población de tortuga mora (Testudo graeca) del Sureste Ibérico. MSc thesis, Universidad Miguel Hernández de Elche, Spain (2007).Linden, H., Wikman, M. Goshawk predation on tetraonids: Availability of prey and diet of the predator in the breeding season. J. Anim. Ecol., 953–968 (1983).Fevold, H. R. & Craighead, J. J. Food requirements of the golden eagle. Auk 75, 312–317 (1958).
Google Scholar 
Collopy, M. W. Food consumption and growth energetics of nestling golden eagles. Wilson Bull. 445–458 (1986).Blanco, J. C., Villafuerte, R. Factores ecológicos que influyen sobre las poblaciones de conejos. Efectos de la enfermedad hemorrágico vírica. TRAGSA, Madrid Spain (1993). More