Philippa Kaur

All methods were performed in accordance with the relevant guidelines and regulations.Study siteThe laboratory experiments on regeneration and wash resistance were conducted at the KCMUCo-PAMVERC Insecticide Testing Facility; while experimental hut study was carried out at Harusini, the facility’s field site located at Mabogini village (S03˚22.764’ E03˚720.793’), adjacent to Lower Moshi rice irrigation scheme in north-eastern Tanzania. The dominant vector at this site is An. arabiensis with moderate level of resistance to pyrethroids conferred by both oxidase and esterase activities32. In this study, pyrethroid-resistant laboratory reared An. gambiae Muleba-Kis mosquitoes were released into the huts for the release-recapture experiment.Test systemsNon-blood fed, 2–5 day old females of susceptible An. gambiae s.s. Kisumu strain and pyrethroid resistant An. gambiae s.s Muleba-Kis strain were used for the evaluation of efficacy in the laboratory (phase I). The Muleba-Kis strain has been colonized for more than 8 years and it is resistant to permethrin with fixed L1014S kdr frequency and metabolic resistance through increased oxidase activity has also been reported21. Only An. gambiae s.s Muleba-Kis were used in release-recapture experiments. The Kisumu strain is fully susceptible to insecticides and free of any detectable insecticide resistance mechanisms. The strain originated from Kisumu, Kenya and has been colonized for many years in laboratory. At the KCMUCo-PAMVERC Moshi insectary, the adult Kisumu strain mosquitoes are reared at a temperature of 24–27 °C, 75 ± 10% relative humidity (RH) and maintained under a dark:light regime of 12:12 h. The Muleba-Kis mosquitoes used for the release-recapture experiments were reared in the field insectary under ambient temperature and relative humidity and treated as previously explained21. The susceptibility status of these colonies is checked every three months using WHO susceptibility test33 and, CDC bottle bioassay test34. The colonies are regularly genotyped for kdr mutations using TaqMan assays35. To maintain the resistance of Muleba-Kis, larvae are frequently selected with alpha-cypermethrin.Regeneration timeTo determine the regeneration time of the insecticide-treated blankets, blankets were cut into 25 × 25 cm pieces and tested before washing and then washed and dried three times consecutively following WHO recommended procedures for LLINs36. The pieces were then re-tested after one, two, three, six and seven days post-washing using WHO cylinders against susceptible An. gambiae s.s (Kisumu).Graphs for 24-h mortality and 60 min knock down (KD) correlating to insecticide bioavailability, as measured by 3 min exposure in cylinder bioassays, were established before and after washing blanket pieces three times consecutively in a day, and tested within a maximum of seven days post-washing. The time in days required to reach initial mortality or 60 min KD plateau is the period required for full regeneration of insecticide-treated blanket.Wash resistanceWHO cylinder bioassays36 were used to assess the wash resistance for the blanket pieces washed 0, 5, 10, 15 and 20 times at the intervals equivalent to the regeneration time. Four pieces cut from 4 permethrin and 4 untreated blankets were used as positive and negative control respectively, against 4 pieces cut from 4 PBO–permethrin blankets.Bioassay proceduresFive, non-blood fed, 2–5 day old An. gambiae Kisumu or An. gambiae Muleba-Kis mosquitoes were exposed for 3 min or 30 min to blanket pieces in WHO cylinder. Bioassays were carried out at 27 ± 2 °C and 75 ± 10% RH. Knock-down was scored after 60 min post-exposure and mortality after 24 h. Fifty mosquitoes (5 mosquitoes per cylinder) were used on each 25 × 25 cm piece of blanket sample. After exposure, the mosquitoes were held for 24 h with access to 10% glucose solution in the paper cups covered with a net material. Mosquitoes exposed to untreated blanket were referred as a negative control.WHO tunnel test methodBlanket pieces which recorded ≤ 80% mortality in cylinder bioassay were tested in the tunnel assay using WHO guidelines. The tunnel was made of an acrylic square cylinder (25 cm in height, 25 cm in width, and 60 cm in length) divided into two sections using a blanket-covered frame fitted into a slot across the tunnel. During the assays a guinea pig was held in a small wooden cage (as a bait) in one of the sections and 50, non-blood fed, female An. gambiae Kisumu or An. gambiae Muleba-Kis aged 5–8 days were released in the other section at dusk and left overnight (13 h) for experimentation at 27 ± 2 °C and 75 ± 10% RH. The blanket surface was deliberately holed (nine 1-cm holes) to allow mosquitoes to contact the blanket material and penetrate to the baited chamber. Treated blankets were tested concurrently together with an untreated blanket. Scoring for the numbers of mosquitoes found alive or dead, fed or unfed, in each section were done in the morning. Mosquitoes found alive were removed and held in paper cups with labels corresponding to each tunnel sections under controlled conditions (25–27 °C and 75–85% RH) and fed on 10% glucose solution to monitor for delayed mortality post exposurely. Outcomes recorded were: mosquito penetration, blood feeding and mortality.Washing of blankets and whole nets for hut trialBlankets and whole nets were separately washed following WHOPES guidelines. In brief, each blanket/net was washed in Savon de Marseilles soap solution (2 g/L) for 10 min: 3 min stirring, 4 min soaking, then another 3 min stirring. This was followed by 2 rinse cycles of the same duration with water only. The water pH was 6 for all washes. The mean water hardness was within the WHOPES limit of ≤ 89 ppm. All nets used in the experimental hut study were cut with holes (4 cm × 4 cm) to simulate the conditions of a torn net. While nets were washed 20 times as per guidelines, blankets were only washed 10 times. To simulate a situation in emergence situations where washing is less frequent due to water scarcity30,31.Experimental hut trial:experimental hut designExperimental hut study was done in Lower Moshi using typical East African experimental huts design as described in the WHOPES35. Huts were constructed with brick walls and featured with cement plaster on the inside and a ceiling board, a metal iron sheet roof, open eaves with window and veranda traps on each side and window traps. Slight modifications from the original structure were made by installing metal eave baffles on two sides. The baffles allow mosquito entry but prevent exits. The window traps were used to collect mosquitoes that tend to exit the huts.Test item labelling, washing and perforatingBoth blankets and LLINs for the trial were distinctively labelled with fabric labels that withstand washes. For wash resistance, the blankets and nets were separately washed according to a protocol adapted from the standard WHO washing procedure36 at the interval equivalent to the regeneration time established in the laboratory for blanket and LLIN respectively. Before testing in the experimental huts, all nets were deliberately holed i.e. 30 holes measuring 4 × 4 cm were made in each net, 9 holes in each of the long side panels, and 6 holes at each short side (head- and foot-side panels) to enhance blood-feeding on the control arm.Test items packagingEach blanket and net were sealed in a plastic bag and then packed in the large plastic container. Each container was labelled for a single treatment to avoid cross contamination between test items.Experimental hut decontaminationA cone assay with 10 susceptible mosquitoes was performed on one wall per hut to rule out any contamination of the wall surface. Only huts with 24 h mortality of susceptible mosquitoes  More

La Sorte, F. A., Johnston, A. & Ault, T. R. Global trends in the frequency and duration of temperature extremes. Clim. Change 166, 1–2 (2021).Article 
ADS 

Google Scholar 
Hansen, J., Sato, M., Ruedy, R., Lo, K. & Medina-Elizade, M. Global temperature change. Proc. Natl. Acad. Sci. U.S.A. 103(39), 14288–14293 (2006).Article 
ADS 
CAS 

Google Scholar 
Borchert, R., Robertson, K., Schwartz, M. D. & Williams-Linera, G. Phenology of temperate trees in tropical climates. Int. J. Biometeorol. 50, 57–65 (2005).Article 
ADS 

Google Scholar 
Misra, G., Sarah, A. & Menzel, A. Ground and satellite phenology in alpine forests are becoming more heterogeneous across higher elevations with warming. Agric. For. Meteorol. 303, 108383 (2021).Article 
ADS 

Google Scholar 
Zuo, Z., Xiao, D. & Qiong, H. Role of the warming trend in global land surface air temperature variations. Sci. China Earth Sci. 6, 866–871 (2021).Article 
ADS 

Google Scholar 
Ling, Y. et al. Assessing the accuracy of forest phenological extraction from sentinel-1 C-band backscatter measurements in deciduous and coniferous forests. Remote Sens. 14(3), 674 (2022).Article 
ADS 

Google Scholar 
Zhang, H., Yuan, W., Liu, S., Dong, W. & Fu, Y. Sensitivity of flowering phenology to changing temperature in China. J. Geophys. Res. Biogeosci. 120(8), 1658–1665 (2015).Article 

Google Scholar 
Cho, J. G. et al. Apple phenology occurs earlier across South Korea with higher temperatures and increased precipitation. Int. J. Biometeorol. 65, 265–276 (2020).Article 

Google Scholar 
Li, C. et al. Response of vegetation phenology to the interaction of temperature and precipitation changes in Qilian mountains. Remote Sens. 14(5), 1248 (2022).Article 
ADS 

Google Scholar 
Berra, E. F. & Gaulton, R. Remote sensing of temperate and boreal forest phenology: A review of progress, challenges and opportunities in the intercomparison of in-situ and satellite phenological metrics. For. Ecol. Manage. 480, 118663 (2021).Article 

Google Scholar 
Zhang, Y. & Li, M. A new method for monitoring start of season (SOS) of forest based on multisource remote sensing. Int. J. Appl. Earth Obs. Geoinf. 104, 102556 (2021).
Google Scholar 
Zhang, X. et al. Monitoring vegetation phenology using MODIS. Remote Sens. Environ. 84(3), 471–475 (2003).Article 
ADS 

Google Scholar 
Thapa, S., Garcia Millan, V. E. & Eklundh, L. Assessing forest phenology: A multi-scale comparison of near-surface (UAV, spectral reflectance sensor, PhenoCam) and Satellite (MODIS, Sentinel-2) remote sensing. Remote Sens. 13, 1597 (2021).Article 
ADS 

Google Scholar 
Bórnez, K., Descals, A., Verger, A. & Peñuelas, J. Land surface phenology from VEGETATION and PROBA-V data: Assessment over deciduous forests. Int. J. Appl. Earth Observ. Geoinf. 84, 101974 (2020).
Google Scholar 
Yu, L., Yan, Z. & Zhang, S. Forest phenology shifts in response to climate change over China–Mongolia–Russia international economic corridor. Forests 11, 757 (2020).Article 

Google Scholar 
Lara, C. et al. Climatic regulation of vegetation phenology in protected areas along Western South America. Remote Sens. 13, 2590 (2021).Article 
ADS 

Google Scholar 
Silveira, E. M. O. et al. Forest phenoclusters for Argentina based on vegetation phenology and climate. Ecol. Appl. 32, 2526 (2022).Article 

Google Scholar 
Tatalovich, Z., Wilson, J. P. & Cockburn, M. A comparison of thiessen polygon, kriging, and spline models of potential UV exposure. Cartogr. Geogr. Inf. Sci. 33, 217–231 (2006).Article 

Google Scholar 
Choubin, B. et al. Spatiotemporal dynamics assessment of snow cover to infer snowline elevation mobility in the mountainous regions. Cold Reg. Sci. Technol. 167, 102870 (2019).Article 

Google Scholar 
Rojas, R., Flexas, J. & Coopman, R. E. Particularities of the highest elevation treeline in the world: Polylepis tarapacana Phil. as a model to study ecophysiological adaptations to extreme environments. Flora 292, 152076 (2022).Article 

Google Scholar 
Du, J. et al. Interacting effects of temperature and precipitation on climatic sensitivity of spring vegetation green-up in arid mountains of China. Agric. For. Meteorol. 269–270, 71–77 (2019).Article 
ADS 

Google Scholar 
Du, J. et al. Daily minimum temperature and precipitation control on spring phenology in arid-mountain ecosystems in China. Int. J. Climatol. 40, 2568–2579 (2020).Article 

Google Scholar 
He, Z. et al. Impacts of recent climate extremes on spring phenology in arid-mountain ecosystems in China. Agric. For. Meteorol. 260–261, 31–40 (2018).Article 
ADS 

Google Scholar 
He, Z. et al. Assessing temperature sensitivity of subalpine shrub phenology in semi-arid mountain regions of China. Agric. For. Meteorol. 213, 42–52 (2015).Article 
ADS 

Google Scholar 
Mu, C., Lu, H., Wang, B., Bao, X. & Cui, W. Short-term effects of harvesting on carbon storage of boreal Larix gmelinii–Carex schmidtii forested wetlands in Daxing’anling, northeast China. For. Ecol. Manage. 293, 140–148 (2013).Article 

Google Scholar 
Hu, T. et al. Effects of fire on soil respiration and its components in a Dahurian larch (Larix gmelinii) forest in northeast China: Implications for forest ecosystem carbon cycling. Geoderma 402, 115273 (2021).Article 
ADS 
CAS 

Google Scholar 
Nyikadzino, B., Chitakira, M. & Muchuru, S. Rainfall and runoff trend analysis in the Limpopo river basin using the Mann Kendall statistic. Phys. Chem. Earth 117, 102870 (2020).Article 

Google Scholar 
Gocic, M. & Trajkovic, S. Analysis of changes in meteorological variables using Mann-Kendall and Sen’s slope estimator statistical tests in Serbia. Glob. Planet. Change 100, 172–182 (2013).Article 
ADS 

Google Scholar 
Fang, Y. et al. Changing contribution rate of heavy rainfall to the rainy season precipitation in Northeast China and its possible causes. Atmos. Res. 197, 437–445 (2017).Article 

Google Scholar 
Piao, S. et al. Changes in satellite-derived vegetation growth trend in temperate and boreal Eurasia from 1982 to 2006. Glob. Change Biol. 17, 3228–3239 (2011).Article 
ADS 

Google Scholar 
Ahas, R., Aasa, A., Menzel, A., Fedotova, V. G. & Scheifinger, H. Changes in European spring phenology. Int. J. Climatol. 22, 1727–1738 (2002).Article 

Google Scholar 
Liang, L., Henebry, G. M., Liu, L., Zhang, X. & Hsu, L. C. Trends in land surface phenology across the conterminous United States (1982–2016) analyzed by NEON domains. Ecol. Appl. 31, e02323 (2021).Article 

Google Scholar 
Fu, Y. H. et al. Decreasing control of precipitation on grassland spring phenology in temperate China. Glob. Ecol. Biogeogr. 30, 490–499 (2020).Article 

Google Scholar 
Aze, T. Unraveling ecological signals from a global warming event of the past. Proc. Natl. Acad. Sci. U.S.A. 119, e2201495119 (2022).Article 

Google Scholar 
Menzel, A., Estrella, N. & Testka, A. Temperature response rates from long-term phenological records. Climate Res. 30, 21–28 (2005).Article 
ADS 

Google Scholar 
Wang, H., Liu, D., Lin, H., Montenegro, A. & Zhu, X. NDVI and vegetation phenology dynamics under the influence of sunshine duration on the Tibetan plateau. Int. J. Climatol. 35, 687–698 (2015).Article 

Google Scholar 
Lesica, P. & Kittelson, P. M. Precipitation and temperature are associated with advanced flowering phenology in a semi-arid grassland. J. Arid Environ. 74, 1013–1017 (2010).Article 
ADS 

Google Scholar 
Shen, M., Piao, S., Cong, N., Zhang, G. & Jassens, I. A. Precipitation impacts on vegetation spring phenology on the Tibetan Plateau. Glob. Change Biol. 21, 3647–3656 (2015).Article 
ADS 

Google Scholar 
Li, Z. et al. Spatio-temporal responses of cropland phenophases to climate change in Northeast China. J. Geog. Sci. 22, 29–45 (2012).Article 
CAS 

Google Scholar 
Badeck, F. W. et al. Responses of spring phenolgy to climate change. New Phytol. 162, 295–309 (2004).Article 

Google Scholar 
Peng, H., Xia, H., Chen, H., Zhi, P. & Xu, Z. Spatial variation characteristics of vegetation phenology and its influencing factors in the subtropical monsoon climate region of southern China. PLoS ONE 16, e0250825 (2021).Article 
CAS 

Google Scholar 
Zhang, J. et al. NIRv and SIF better estimate phenology than NDVI and EVI: Effects of spring and autumn phenology on ecosystem production of planted forests. Agric. For. Meteorol. 315, 108819 (2022).Article 
ADS 

Google Scholar 
Yu, X., Zhuang, D., Hou, X. & Chen, H. Forest phenological patterns of Northeast China inferred from MODIS data. J. Geog. Sci. 15, 239–246 (2005).Article 

Google Scholar 
Chen, X. & Xu, L. Phenological responses of Ulmus pumila (Siberian Elm) to climate change in the temperate zone of China. Int. J. Biometeorol. 56, 695–706 (2012).Article 
ADS 

Google Scholar 
Ma, X., Bai, H., He, Y. & Li, S. The vegetation RSP of Qinling Mountains based on the NDVI and the response of temperature to it. Appl. Mech. Mater. 700, 394–399 (2014).Article 

Google Scholar  More

Moberg, F. & Folke, C. Ecological goods and services of coral reef ecosystems. Ecol. Econ. 29, 215–233 (1999).Article 

Google Scholar 
Woodhead, A. J., Hicks, C. C., Norström, A. V., Williams, G. J. & Graham, N. A. J. Coral reef ecosystem services in the Anthropocene. Funct. Ecol. 33, 1023–1034 (2019).
Google Scholar 
Hughes, T. P. et al. Coral reefs in the Anthropocene. Nature 546, 82 (2017).Article 
ADS 
CAS 

Google Scholar 
Hughes, T. P., Kerry, J. T. & Simpson, T. Large-scale bleaching of corals on the Great Barrier Reef. Ecology 99, 501 (2017).Article 

Google Scholar 
Anthony, K. R. N., Kline, D. I., Diaz-Pulido, G., Dove, S. & Hoegh-Guldberg, O. Ocean acidification causes bleaching and productivity loss in coral reef builders. PNAS 105, 17442–17446 (2008).Article 
ADS 
CAS 

Google Scholar 
Courtial, L., Roberty, S., Shick, J. M., Houlbrèque, F. & Ferrier-Pagès, C. Interactive effects of ultraviolet radiation and thermal stress on two reef-building corals. Limnol. Oceanogr. 62, 1000–1013 (2017).Article 
ADS 

Google Scholar 
Jessen, C. et al. In-situ effects of eutrophication and overfishing on physiology and bacterial diversity of the Red Sea Coral Acropora hemprichii. PLoS ONE 8, e62091 (2013).Article 
ADS 
CAS 

Google Scholar 
Jessen, C., Roder, C., Villa Lizcano, J. F., Voolstra, C. R. & Wild, C. In-situ effects of simulated overfishing and eutrophication on benthic coral reef algae growth, succession, and composition in the Central Red Sea. PLoS ONE 8, e66992 (2013).Article 
ADS 
CAS 

Google Scholar 
Fabricius, K. E. Effects of terrestrial runoff on the ecology of corals and coral reefs: Review and synthesis. Mar. Pollut. Bull. 50, 125–146 (2005).Article 
CAS 

Google Scholar 
Hughes, T. P. et al. Climate change, human impacts, and the resilience of coral reefs. Science 301, 929–933 (2003).Article 
ADS 
CAS 

Google Scholar 
Fabricius, K. E., Cséke, S., Humphrey, C. & De’ath, G. Does trophic status enhance or reduce the thermal tolerance of scleractinian corals? A review, experiment and conceptual framework. PLoS ONE 8, e54399 (2013).Article 
ADS 
CAS 

Google Scholar 
McLachlan, R. H., Price, J. T., Solomon, S. L. & Grottoli, A. G. Thirty years of coral heat-stress experiments: A review of methods. Coral Reefs 39, 885–902 (2020).Article 

Google Scholar 
Fabricius, K. E. Factors determining the resilience of coral reefs to eutrophication: A review and conceptual model. In Coral Reefs: An Ecosystem in Transition (eds Dubinsky, Z. & Stambler, N.) (Springer, 2011).
Google Scholar 
Tilstra, A. et al. Light induced intraspecific variability in response to thermal stress in the hard coral Stylophora pistillata. PeerJ. https://doi.org/10.7717/PEERJ.3802/ (2017).Article 

Google Scholar 
Connolly, S. R., Lopez-Yglesias, M. A. & Anthony, K. R. N. Food availability promotes rapid recovery from thermal stress in a scleractinian coral. Coral Reefs 31, 951–960 (2012).Article 
ADS 

Google Scholar 
Coles, S. L. & Brown, B. E. Coral bleaching—Capacity for acclimatization and adaptation. Adv. Mar. Biol. 46, 183 (2003).Article 
CAS 

Google Scholar 
Rosenberg, E., Koren, O., Reshef, L., Efrony, R. & Zilber-Rosenberg, I. The role of microorganisms in coral health, disease and evolution. Nat. Rev. Microbiol. 5, 355–362 (2007).Article 
CAS 

Google Scholar 
Szmant, A. M. Nutrient enrichment on coral reefs: Is it a major cause of coral reef decline? Estuaries 25, 743–766 (2002).Article 
CAS 

Google Scholar 
Atkinson, M. J., Carlson, B. & Crow, G. L. Coral growth in high-nutrient, low-pH seawater: A case study of corals cultured at the Waikiki Aquarium, Honolulu, Hawaii. Coral Reefs 14, 215–223 (1995).Article 
ADS 

Google Scholar 
Bongiorni, L., Shafir, S., Angel, D. & Rinkevich, B. Survival, growth and gonad development of two hermatypic corals subjected to in situ fish-farm nutrient enrichment. Mar. Ecol. Prog. Ser. 253, 137–144 (2003).Article 
ADS 

Google Scholar 
Grigg, R. W. Coral reefs in an urban embayment in Hawaii: A complex case history controlled by natural and anthropogenic stress. Coral Reefs 14, 253–266 (1995).Article 
ADS 

Google Scholar 
Fabricius, K. E. & De’ath, G. Identifying ecological change and its causes: A case study on coral reefs. Ecol. Appl. 14, 1448–1465 (2004).Article 

Google Scholar 
Ferrier-Pagès, C., Gattuso, J. P., Dallot, S. & Jaubert, J. Effect of nutrient enrichment on growth and photosynthesis of the zooxanthellate coral Stylophora pistillata. Coral Reefs 19, 103–113 (2000).Article 

Google Scholar 
Rosset, S., Wiedenmann, J., Reed, A. J. & D’Angelo, C. Phosphate deficiency promotes coral bleaching and is reflected by the ultrastructure of symbiotic dinoflagellates. Mar. Pollut. Bull. 118, 180–187 (2017).Article 
CAS 

Google Scholar 
Ban, S. S., Graham, N. A. J. & Connolly, S. R. Evidence for multiple stressor interactions and effects on coral reefs. Glob. Change Biol. 20, 681–697 (2014).Article 
ADS 

Google Scholar 
Wiedenmann, J. et al. Nutrient enrichment can increase the susceptibility of reef corals to bleaching. Nat. Clim. Change 3, 160–164 (2012).Article 
ADS 

Google Scholar 
Rädecker, N. et al. Heat stress destabilizes symbiotic nutrient cycling in corals. PNAS. https://doi.org/10.1073/pnas.2022653118 (2021).Article 

Google Scholar 
LaJeunesse, T. C. et al. Systematic revision of symbiodiniaceae highlights the antiquity and diversity of coral endosymbionts. Curr. Biol. 28, 2570–2580 (2018).Article 
CAS 

Google Scholar 
Falkowski, P. G., Dubinsky, Z., Muscatine, L. & McCloskey, L. Population control in symbiotic corals—Ammonium ions and organic materials maintain the density of zooxanthellae. Bioscience 43, 606–611 (1993).Article 

Google Scholar 
Muscatine, L. & Pool, R. R. Regulation of numbers of intracellular algae. Proc. R. Soc. Lond. Ser. B Biol. Sci. 204, 131–139 (1979).ADS 
CAS 

Google Scholar 
Muller-Parker, G., D’Elia, C. F. & Cook, C. B. Interactions between corals and their symbiotic algae. Coral Reefs Anthr. https://doi.org/10.1007/978-94-017-7249-5_5 (2015).Article 

Google Scholar 
Rädecker, N., Pogoreutz, C., Voolstra, C. R., Wiedenmann, J. & Wild, C. Nitrogen cycling in corals: The key to understanding holobiont functioning? Trends Microbiol. 23, 490–497 (2015).Article 

Google Scholar 
Steinberg, R. K., Dafforn, K. A., Ainsworth, T. & Johnston, E. L. Know thy anemone: A review of threats to octocorals and anemones and opportunities for their restoration. Front. Mar. Sci. 7, 590 (2020).Article 

Google Scholar 
Inoue, S., Kayanne, H., Yamamoto, S. & Kurihara, H. Spatial community shift from hard to soft corals in acidified water. Nat. Clim. Change 3, 683–687 (2013).Article 
ADS 
CAS 

Google Scholar 
Wild, C. & Naumann, M. S. Effect of active water movement on energy and nutrient acquisition in coral reef-associated benthic organisms. PNAS 110, 8767–8768 (2013).Article 
ADS 
CAS 

Google Scholar 
Fox, H. E., Pet, J. S., Dahuri, R. & Caldwell, R. L. Recovery in rubble fields: Long-term impacts of blast fishing. Mar. Pollut. Bull. 46, 1024–1031 (2003).Article 
CAS 

Google Scholar 
Benayahu, Y. & Loya, Y. Settlement and recruitment of a soft coral: Why is Xenia macrospiculata a successful colonizer? Bull. Mar. Sci. 36, 177–188 (1985).
Google Scholar 
Norström, A. V., Nyström, M., Lokrantz, J. & Folke, C. Alternative states on coral reefs: Beyond coral-macroalgal phase shifts. Mar. Ecol. Prog. Ser. 376, 293–306 (2009).Article 
ADS 

Google Scholar 
Reverter, M., Helber, S. B., Rohde, S., De Goeij, J. M. & Schupp, P. J. Coral reef benthic community changes in the Anthropocene: Biogeographic heterogeneity, overlooked configurations, and methodology. Glob. Change Biol. 28, 1956–1971 (2022).Article 

Google Scholar 
Karcher, D. B. et al. Nitrogen eutrophication particularly promotes turf algae in coral reefs of the central Red Sea. PeerJ 2020, 1–25 (2020).
Google Scholar 
El-Khaled, Y. C. et al. Nitrogen fixation and denitrification activity differ between coral- and algae-dominated Red Sea reefs. Sci. Rep. 11, 1–15 (2021).Article 

Google Scholar 
Ruiz-Allais, J. P., Benayahu, Y. & Lasso-Alcalá, O. M. The invasive octocoral Unomia stolonifera (Alcyonacea, Xeniidae) is dominating the benthos in the Southeastern Caribbean Sea. Mem. la Fund La Salle Ciencias Nat. 79, 63–80 (2021).
Google Scholar 
Ruiz Allais, J. P., Amaro, M. E., McFadden, C. S., Halász, A. & Benayahu, Y. The first incidence of an alien soft coral of the family Xeniidae in the Caribbean, an invasion in eastern Venezuelan coral communities. Coral Reefs 33, 287 (2014).Article 
ADS 

Google Scholar 
Baum, G., Januar, I., Ferse, S. C. A., Wild, C. & Kunzmann, A. Abundance and physiology of dominant soft corals linked to water quality in Jakarta Bay, Indonesia. PeerJ 2016, 1–29 (2016).
Google Scholar 
Menezes, N. M. et al. New non-native ornamental octocorals threatening a South-west Atlantic reef. J. Mar. Biol. Assoc. U.K. https://doi.org/10.1017/S0025315421000849 (2022).Article 

Google Scholar 
Mantelatto, M. C., da Silva, A. G., dos Louzada, T. S., McFadden, C. S. & Creed, J. C. Invasion of aquarium origin soft corals on a tropical rocky reef in the southwest Atlantic. Brazil. Mar. Pollut. Bull. 130, 84–94 (2018).Article 
CAS 

Google Scholar 
Simancas-Giraldo, S. M. et al. Photosynthesis and respiration of the soft coral Xenia umbellata respond to warming but not to organic carbon eutrophication. PeerJ 9, e11663 (2021).Article 

Google Scholar 
Vollstedt, S., Xiang, N., Simancas-Giraldo, S. M. & Wild, C. Organic eutrophication increases resistance of the pulsating soft coral Xenia umbellata to warming. PeerJ 2020, 1–16 (2020).
Google Scholar 
Thobor, B. et al. The pulsating soft coral Xenia umbellata shows high resistance to warming when nitrate concentrations are low. Sci. Rep. https://doi.org/10.1038/s41598-022-21110-w (2022).Article 

Google Scholar 
Costa, O. S., Leão, Z. M. A. N., Nimmo, M. & Attrill, M. J. Nutrification impacts on coral reefs from northern Bahia, Brazil. Hydrobiologia 440, 307–315 (2000).Article 
CAS 

Google Scholar 
Fleury, B. G., Coll, J. C., Tentori, E., Duquesne, S. & Figueiredo, L. Effect of nutrient enrichment on the complementary (secondary) metabolite composition of the soft coral Sarcophyton ebrenbergi (Cnidaria: Octocorallia: Alcyonaceae) of the Great Barrier Reef. Mar. Biol. 136, 63–68 (2000).Article 
CAS 

Google Scholar 
Bednarz, V. N., Naumann, M. S., Niggl, W. & Wild, C. Inorganic nutrient availability affects organic matter fluxes and metabolic activity in the soft coral genus Xenia. J. Exp. Biol. 215, 3672–3679 (2012).CAS 

Google Scholar 
Bruno, J. F., Petes, L. E., Harvell, C. D. & Hettinger, A. Nutrient enrichment can increase the severity of coral diseases. Ecol. Lett. 6, 1056–1061 (2003).Article 

Google Scholar 
Ezzat, L., Maguer, J.-F.F., Grover, R. & Ferrier-Pagès, C. Limited phosphorus availability is the Achilles heel of tropical reef corals in a warming ocean. Sci. Rep. 6, 1–11 (2016).Article 

Google Scholar 
Tanaka, Y., Grottoli, A. G., Matsui, Y., Suzuki, A. & Sakai, K. Effects of nitrate and phosphate availability on the tissues and carbonate skeleton of scleractinian corals. Mar. Ecol. Prog. Ser. 570, 101–112 (2017).Article 
ADS 
CAS 

Google Scholar 
Liu, G., Strong, A. E., Skirving, W. & Arzayus, L. F. Overview of NOAA coral reef watch program’s near-real time satellite global coral bleaching monitoring activities. In Proc. 10th International Coral Reef Symposium, 1783–1793 (2006).Bellworthy, J. & Fine, M. Beyond peak summer temperatures, branching corals in the Gulf of Aqaba are resilient to thermal stress but sensitive to high light. Coral Reefs 36, 1071–1082 (2017).Article 
ADS 

Google Scholar 
Rex, A., Montebon, F. & Yap, H. T. Metabolic responses of the scleractinian coral Porites cylindrica Dana to water motion. I. Oxygen flux studies. J. Exp. Mar. Biol. Ecol. 186, 33–52 (1995).Article 

Google Scholar 
Long, M. H., Berg, P., de Beer, D. & Zieman, J. C. In situ coral reef oxygen metabolism: An eddy correlation study. PLoS ONE 8, e58581 (2013).Article 
ADS 
CAS 

Google Scholar 
Fabricius, K. E. & Klumpp, D. W. Widespread mixotrophy in reef-inhabiting soft corals: The influence of depth, and colony expansion and contraction on photosynthesis. Mar. Ecol. Prog. Ser. 125, 195–204 (1995).Article 
ADS 

Google Scholar 
Bradford, M. M. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal. Biochem. 72, 248–254 (1976).Article 
CAS 

Google Scholar 
Raimonet, M., Guillou, G., Mornet, F. & Richard, P. Macroalgae δ15N values in well-mixed estuaries: Indicator of anthropogenic nitrogen input or macroalgae metabolism? Estuar. Coast. Shelf Sci. 119, 126–138 (2013).Article 
ADS 
CAS 

Google Scholar 
Furla, P., Galgani, I., Durand, I. & Allemand, D. Sources and mechanisms of inorganic carbon transport for coral calcification and photosynthesis. J. Exp. Biol. 203, 3445–3457 (2000).Article 
CAS 

Google Scholar 
Hughes, A. D., Grottoli, A. G., Pease, T. K. & Matsui, Y. Acquisition and assimilation of carbon in non-bleached and bleached corals. Mar. Ecol. Prog. Ser. 420, 91–101 (2010).Article 
ADS 
CAS 

Google Scholar 
Rau, G. H., Takahashi, T. & Des Marais, D. J. Latitudinal variations in plankton delta C-13—Implications for CO2 and productivity in past oceans. Nature 341, 516–518 (1989).Article 
ADS 
CAS 

Google Scholar 
McMahon, K. W., Hamady, L. L. & Thorrold, S. R. A review of ecogeochemistry approaches to estimating movements of marine animals. Limnol. Oceanogr. 58, 697–714 (2013).Article 
ADS 
CAS 

Google Scholar 
Muscatine, L., Porter, J. W. & Kaplan, I. R. Resource partitioning by reef corals as determined from stable isotope composition. Mar. Biol. 100, 185–193 (1989).Article 

Google Scholar 
Swart, P. K. et al. The isotopic composition of respired carbon dioxide in scleractinian corals: Implications for cycling of organic carbon in corals. Geochim. Cosmochim. Acta 69, 1495–1509 (2005).Article 
ADS 
CAS 

Google Scholar 
Rodrigues, L. J. & Grottoli, A. G. Calcification rate and the stable carbon, oxygen, and nitrogen isotopes in the skeleton, host tissue, and zooxanthellae of bleached and recovering Hawaiian corals. Geochim. Cosmochim. Acta 70, 2781–2789 (2006).Article 
ADS 
CAS 

Google Scholar 
Grottoli, A. G. & Rodrigues, L. J. Bleached Porites compressa and Montipora capitata corals catabolize δ13C-enriched lipids. Coral Reefs 30, 687–692 (2011).Article 
ADS 

Google Scholar 
Levas, S. J., Grottoli, A. G., Hughes, A., Osburn, C. L. & Matsui, Y. Physiological and biogeochemical traits of bleaching and recovery in the mounding species of coral porites lobata: Implications for resilience in mounding corals. PLoS ONE 8, 32–35 (2013).Article 

Google Scholar 
Schoepf, V. et al. Annual coral bleaching and the long-term recovery capacity of coral. Proc. R. Soc. B Biol. Sci. 282, 20151887 (2015).Article 

Google Scholar 
Lesser, M. P. et al. Nitrogen fixation by symbiotic cyanobacteria provides a source of nitrogen for the scleractinian coral Montastraea cavernosa. Mar. Ecol. Prog. Ser. 346, 143–152 (2007).Article 
ADS 
CAS 

Google Scholar 
Carpenter, E. J., Harvey, H. R., Brian, F. & Capone, D. G. Biogeochemical tracers of the marine cyanobacterium Trichodesmium. Deep Sea Res. I Oceanogr. Res. Pap. 44, 27–38 (1997).Article 
ADS 
CAS 

Google Scholar 
Lachs, L. et al. Effects of tourism-derived sewage on coral reefs: Isotopic assessments identify effective bioindicators. Mar. Pollut. Bull. 148, 85–96 (2019).Article 
CAS 

Google Scholar 
Kürten, B. et al. Influence of environmental gradients on C and N stable isotope ratios in coral reef biota of the Red Sea, Saudi Arabia. J. Sea Res. 85, 379–394 (2014).Article 
ADS 

Google Scholar 
Core Team, R. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2020).Wickham, H. et al. Welcome to the Tidyverse. J. Open Source Softw. 4, 1686 (2019).Article 
ADS 

Google Scholar 
Kassambara, A. ggpubr: ‘ggplot2’ Based Publication Ready Plots. R Package Version 0.4.0 (2020).Kassambara, A. rstatix: Pipe-Friendly Framework for Basic Statistical Tests. R Package Version 0.7.0 (2021).Contreras-Silva, A. I. et al. A meta-analysis to assess long-term spatiotemporal changes of benthic coral and macroalgae cover in the Mexican Caribbean. Sci. Rep. 10, 1–12 (2020).Article 

Google Scholar 
Ledlie, M. H. et al. Phase shifts and the role of herbivory in the resilience of coral reefs. Coral Reefs 26, 641–653 (2007).Article 
ADS 

Google Scholar 
Kuffner, I. B. & Toth, L. T. A geological perspective on the degradation and conservation of western Atlantic coral reefs. Conserv. Biol. 30, 706–715 (2016).Article 

Google Scholar 
Hughes, T. P. Catastrophes, phase shifts, and large-scale degradation of a Caribbean Coral Reef. Science 265, 1547–1551 (1994).Article 
ADS 
CAS 

Google Scholar 
de Bakker, D. M., Meesters, E. H., Bak, R. P. M., Nieuwland, G. & van Duyl, F. C. Long-term shifts in coral communities on shallow to deep reef slopes of Curaçao and Bonaire: Are there any winners? Front. Mar. Sci. 3, 247 (2016).Article 

Google Scholar 
Mergner, H. & Svoboda, A. Productivity and seasonal changes in selected reef areas in the Gulf of Aqaba (Red Sea). Helgoländer Meeresun. 30, 383–399 (1977).Article 

Google Scholar 
Schlichter, D., Svoboda, A. & Kremer, B. P. Functional autotrophy of Heteroxenia fuscescens (Anthozoa: Alcyonaria): Carbon assimilation and translocation of photosynthates from symbionts to host. Mar. Biol. 78, 29–38 (1983).Article 
CAS 

Google Scholar 
Al-Sofyani, A. A. & Niaz, G. R. A comparative study of the components of the hard coral Seriatopora hystrix and the soft coral Xenia umbellata along the Jeddah coast, Saudi Arabia. Rev. Biol. Mar. Oceanogr. 42, 207–219 (2007).Article 

Google Scholar 
McCloskey, L. R., Wethey, D. S. & Porter, J. W. Measurement and interpretation of photosynthesis and respiration in reef corals. In Coral Reefs: Research Methods (eds Stoddart, D. R. & Johannes, R. E.) 379–396 (United Nations Educational, Scientific and Cultural Organization, 1978).
Google Scholar 
Baker, D. M., Freeman, C. J., Wong, J. C. Y., Fogel, M. L. & Knowlton, N. Climate change promotes parasitism in a coral symbiosis. ISME J. 12, 921–930 (2018).Article 
CAS 

Google Scholar 
Hoegh-Guldberg, O. & Smith, G. J. The effect of sudden changes in temperature, light and salinity on the population density and export of zooxanthellae from the reef corals Stylophora pistillata Esper and Seriatopora hystrix Dana. J. Exp. Mar. Biol. Ecol. 129, 279–303 (1989).Article 

Google Scholar 
Iglesias-Prieto, R., Matta, J. L., Robins, W. A. & Trench, R. K. Photosynthetic response to elevated temperature in the symbiotic dinoflagellate Symbiodinium microadriaticum in culture. Proc. Natl. Acad. Sci. 89, 10302–10305 (1992).Article 
ADS 
CAS 

Google Scholar 
Béraud, E., Gevaert, F., Rottier, C. & Ferrier-Pagès, C. The response of the scleractinian coral Turbinaria reniformis to thermal stress depends on the nitrogen status of the coral holobiont. J. Exp. Biol. 216, 2665–2674 (2013).
Google Scholar 
Kremien, M., Shavit, U., Mass, T. & Genin, A. Benefit of pulsation in soft corals. Proc. Natl. Acad. Sci. U.S.A. 110, 8978–8983 (2013).Article 
ADS 
CAS 

Google Scholar 
Grover, R. et al. Coral uptake of inorganic phosphorus and nitrogen negatively affected by simultaneous changes in temperature and pH. PLoS ONE 6, 1–10 (2011).
Google Scholar 
Cardini, U. et al. Microbial dinitrogen fixation in coral holobionts exposed to thermal stress and bleaching. Environ. Microbiol. 18, 2620–2633 (2016).Article 
CAS 

Google Scholar 
Cardini, U. et al. Functional significance of dinitrogen fixation in sustaining coral productivity under oligotrophic conditions. Proc. R. Soc. B Biol. Sci. 282, 20152257 (2015).Article 

Google Scholar 
Santos, H. F. et al. Climate change affects key nitrogen-fixing bacterial populations on coral reefs. ISME J. 8, 2272–2279 (2014).Article 

Google Scholar 
Tilstra, A. et al. Relative diazotroph abundance in symbiotic red sea corals decreases with water depth. Front. Mar. Sci. 6, 372 (2019).Article 

Google Scholar 
Klinke, A. et al. Impact of phosphate enrichment on the susceptibility of the pulsating soft coral Xenia umbellata to ocean warming. Front. Mar. Sci. 9, 1026321 (2022).Article 

Google Scholar 
Rädecker, N. et al. Heat stress reduces the contribution of diazotrophs to coral holobiont nitrogen cycling. ISME J. https://doi.org/10.1038/s41396-021-01158-8 (2021).Article 

Google Scholar 
Swart, P. K., Saied, A. & Lamb, K. Temporal and spatial variation in the δ15N and δ13C of coral tissue and zooxanthellae in Montastraea faveolata collected from the Florida reef tract. Limnol. Oceanogr. 50, 1049–1058 (2005).Article 
ADS 
CAS 

Google Scholar 
Grottoli, A. G., Tchernov, D. & Winters, G. Physiological and biogeochemical responses of super-corals to thermal stress from the Northern Gulf of Aqaba, Red Sea. Front. Mar. Sci. 4, 215 (2017).Article 

Google Scholar 
Dubinsky, Z. & Stambler, N. Marine pollution and coral reefs. Glob. Change Biol. 2, 511–526 (1996).Article 
ADS 

Google Scholar 
Loya, Y., Lubinevsky, H., Rosenfeld, M. & Kramarsky-Winter, E. Nutrient enrichment caused by in situ fish farms at Eilat, Red Sea is detrimental to coral reproduction. Mar. Pollut. Bull. 49, 344–353 (2004).Article 
CAS 

Google Scholar 
Costa, O. S., Nimmo, M. & Attrill, M. J. Coastal nutrification in Brazil: A review of the role of nutrient excess on coral reef demise. J. S. Am. Earth Sci. 25, 257–270 (2008).Article 

Google Scholar 
Tait, D. R. et al. The influence of groundwater inputs and age on nutrient dynamics in a coral reef lagoon. Mar. Chem. 166, 36–47 (2014).Article 
CAS 

Google Scholar 
Guan, Y., Hohn, S., Wild, C. & Merico, A. Vulnerability of global coral reef habitat suitability to ocean warming, acidification and eutrophication. Glob. Change Biol. 26, 5646–5660 (2020).Article 
ADS 

Google Scholar 
Hall, E. R. et al. Eutrophication may compromise the resilience of the Red Sea coral Stylophora pistillata to global change. Mar. Pollut. Bull. 131, 701–711 (2018).Article 
CAS 

Google Scholar 
Naumann, M. S. et al. Organic matter release by dominant hermatypic corals of the Northern Red Sea. Coral Reefs 29, 649–659 (2010).Article 
ADS 

Google Scholar 
Wild, C. et al. Coral mucus functions as an energy carrier and particle trap in the reef ecosystem. Nature 428, 66–70 (2004).Article 
ADS 
CAS 

Google Scholar  More

Jordan, D. S. A classification of fishes including families and genera as far as know. Stanford University Publications. Bio. Sci. 3, 79–243. https://doi.org/10.5962/bhl.title.161386 (1923).Article 

Google Scholar 
Akihito, et al. Evolutionary aspects of gobioid fishes based on an analysis of mitochondrial cytochrome b genes. Gene 259, 5–15 (2000).Article 
CAS 

Google Scholar 
Wang, H.-Y., Tsai, M.-P., Dean, J. & Lee, S.-C. Molecular phylogeny of gobioid Wshes (Perciformes: Gobioidei) based on mitochondrial 12S rRNA sequences. Mol. Phylogenet. Evol. 20, 390–408. https://doi.org/10.1016/j.ympev.2005.05.004 (2001).Article 
CAS 

Google Scholar 
Nelson, J. S., Grande, T. C. & Wilson, M. V. Fishes of the World (Wiley, 2016).Book 

Google Scholar 
Fricke, R., Eschmeyer, W. N. & Van der Laan, R. Eschmeyer’s Catalog of fishes: Genera, Species, references. (http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp) (Accessed 15 June 2022).Guimarães-Costa, A. et al. Molecular evidence of two new species of Eleotris (Gobiiformes: Eleotridae) in the western Atlantic. Mol. Phylogenet. Evol. 98, 52–56. https://doi.org/10.1016/j.ympev.2016.01.014 (2016).Article 

Google Scholar 
Thacker, C. E. & Hardman, M. A. Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei). Mol. Phylogenet. Evol. 37, 858–887. https://doi.org/10.1016/j.ympev.2005.05.004 (2005).Article 
CAS 

Google Scholar 
Nordlie, F. G. Life-history characteristics of eleotrid fishes of the western hemisphere, and perils of life in a vanishing environment. Rev. Fish Biol. Fisher. 22(1), 189–224. https://doi.org/10.1007/s11160-011-9229-3 (2012).Article 

Google Scholar 
Berra, T. M. Freshwater Fish Distribution (Academic Press, 2001).
Google Scholar 
Graham, J. B. Air-Breathing Fishes: Evolution, Diversity, and Adaptation (Academic Press, 1997).Book 

Google Scholar 
Thacker, C. E. Phylogeny of Gobioidea and its placement within Acanthomorpha, with a new classification and investigation of diversification and character evolution. Copeia 1, 93–104. https://doi.org/10.1643/CI-08-004 (2009).Article 

Google Scholar 
Chakrabarty, P., Davis, M. P. & Sparks, J. S. The first record of a trans-oceanic sister-group relationship between obligate vertebrate troglobites. PLoS One 7, e44083. https://doi.org/10.1371/journal.pone.0044083 (2012).Article 
ADS 
CAS 

Google Scholar 
Agorreta, A. et al. Molecular phylogenetics of Gobioidei and phylogenetic placement of European gobies. Mol. Phylogenet. Evol. 69, 619–633. https://doi.org/10.1016/j.ympev.2013.07.017 (2013).Article 

Google Scholar 
McCraney, W. T., Thacker, C. E. & Alfaro, M. E. Supermatrix phylogeny resolves goby lineages and reveals unstable root of Gobiaria. Mol. Phylogenet. Evol. 151, 106862. https://doi.org/10.1016/j.ympev.2020.106862 (2020).Article 

Google Scholar 
Karl, S. A. & Avise, J. C. Balancing selection at allozyme loci in oysters: Implications from nuclear RFLPs. Science 256, 100. https://doi.org/10.1126/science.1348870 (1992).Article 
ADS 
CAS 

Google Scholar 
Hey, J. & Machado, C. A. The study of structured populations—New hope for a difficult and divided science. Nat. Rev. Genet. 4, 535–543. https://doi.org/10.1038/nrg1112 (2003).Article 
CAS 

Google Scholar 
Castroviejo-Fisher, S., Guayasamin, J. M., Gonzalez-Voyer, A. & Vilà, C. Neotropical diversification seen through glassfrogs. J. Biogeogr. 41, 66–80. https://doi.org/10.1111/jbi.12208 (2014).Article 

Google Scholar 
Dornburg, A., Townsend, J. P., Friedman, M. & Near, T. J. Phylogenetic informativeness reconciles ray-finned fish molecular divergence times. BMC Evol. Biol. 14, 169. https://doi.org/10.1186/s12862-014-0169-0 (2014).Article 

Google Scholar 
Hundt, P. J., Iglésias, S. P., Hoey, A. S. & Simons, A. M. A multilocus molecular phylogeny of combtooth blennies (Percomorpha: Blennioidei: Blenniidae): Multiple invasions of intertidal habitats. Mol. Phylogenet. Evol. 70, 47–56. https://doi.org/10.1016/j.ympev.2013.09.001 (2014).Article 

Google Scholar 
Olave, M., Avila, L. J., Sites, J. W. & Morando, M. Multilocus phylogeny of the widely distributed South American lizard clade Eulaemus (Liolaemini, Liolaemus). Zool. Scr. 43, 323–337. https://doi.org/10.1111/zsc.12053 (2014).Article 

Google Scholar 
Meyer, B. S., Matschiner, M. & Salzburger, W. A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach. Mol. Phylogenet. Evol. 83, 56–71. https://doi.org/10.1016/j.ympev.2014.10.009 (2015).Article 

Google Scholar 
Jønsson, K. A. et al. A supermatrix phylogeny of corvoid passerine birds (Aves: Corvides). Mol. Phylogenet. Evol. 94, 87–94. https://doi.org/10.1016/j.ympev.2015.08.020 (2016).Article 

Google Scholar 
Li, H. & Durbin, R. Inference of human population history from individual whole-genome sequences. Nature 475(7357), 493–496. https://doi.org/10.1038/nature10231 (2011).Article 
CAS 

Google Scholar 
Frantz, R. S. X-efficiency: Theory, Evidence and Applications Vol. 2 (Springer Science & Business Media, 2013).
Google Scholar 
Bessa-Silva, A. et al. The roles of vicariance and dispersal in the differentiation of two species of the Rhinella marina species complex. Mol. Phylogenet. Evol. 145, 106723. https://doi.org/10.1016/j.ympev.2019.106723 (2020).Article 

Google Scholar 
Leutenegger, W. Maternal-fetal weight relationships in primates. Folia Primatol. 20(4), 280–293. https://doi.org/10.1159/000155580 (1973).Article 
CAS 

Google Scholar 
Yeh, J. The effect of miniaturized body size on skeletal morphology in frogs. Evolution 56(3), 628–641. https://doi.org/10.1111/j.0014-3820.2002.tb01372.x (2002).Article 

Google Scholar 
Daza, J. D. et al. An enigmatic miniaturized and attenuate whole lizard from the Mid-Cretaceous amber of Myanmar. Breviora 563(1), 1–18. https://doi.org/10.3099/MCZ49.1 (2018).Article 

Google Scholar 
Hanken, J. & Wake, D. B. Miniaturization of body size: Organismal consequences and evolutionary significance. Annu. Rev. Ecol. Evol. Syst. 24(1), 501–519. https://doi.org/10.1146/annurev.es.24.110193.002441 (1993).Article 

Google Scholar 
Britz, R. & Conway, K. W. Osteology of Paedocypris, a miniature and highly developmentally truncated fish (Teleostei: Ostariophysi: Cyprinidae). J. Morphol. 270(4), 389–412. https://doi.org/10.1002/jmor.10698 (2009).Article 
CAS 

Google Scholar 
Britz, R., Conway, K. W. & Ruber, L. Spectacular morphological novelty in a miniature cyprinid fish, Danionella dracula n. sp.. Proc. R. Soc. Lond. 276(1665), 2179–2186. https://doi.org/10.1098/rspb.2009.0141 (2009).Article 

Google Scholar 
Weitzman, S. H. & Vari, R. P. Miniaturization in South American freshwater fishes; an overview and discussion. Proc. Biol. Soc. Wash. 101(2), 444–465 (1988).
Google Scholar 
Toledo-Piza, M., Mattox, G. M. & Britz, R. Priocharax nanus, a new miniature characid from the rio Negro, Amazon basin (Ostariophysi: Characiformes), with an updated list of miniature Neotropical freshwater fishes. Neotrop. Ichthyol. 12(2), 229–246. https://doi.org/10.1590/1982-0224-20130171 (2014).Article 

Google Scholar 
Caires, R. A. & Figueiredo, J. L. Review of the genus Microphilypnus Myers, 1927 (Teleostei: Gobioidei: Eleotridae) from the lower Amazon basin, with description of one new species. Zootaxa 3036, 39–57. https://doi.org/10.11646/zootaxa.3036.1.3 (2011).Article 

Google Scholar 
Caires, R. A. Microphilypnus tapajosensis, a new species of eleotridid from the Tapajós basin, Brazil (Gobioidei: Eleotrididae). Ichthyol. Explor. Freshw. 23, 155–160 (2013).
Google Scholar 
Caires, R. A. & Guimarães-Costa, A. Family Eleotridae. In Field Guide to Amazonian Fishes (eds van Sleen, P. & Albert, J.) 388–391 (Princeton University Press, 2017).
Google Scholar 
Caires, R. A. & Toledo-Piza, M. A New species of miniature fish of the genus Microphilypnus (Gobioidei: Eleotridae) from the upper Rio Negro Basin, Amazonas Brazil. Copeia 106(1), 49–55. https://doi.org/10.1643/CI-17-634 (2018).Article 

Google Scholar 
Roberts, T.R. Leptophilypnion, a new genus with two new species of tiny central Amazonian gobioid fishes (Teleostei, Eleotridae). Aqua (2013).Gould, R. E. & Delevoryas, T. The biology of Glossopteris: Evidence from petrified seed-bearing and pollen-bearing organs. Alcheringa 1(4), 387–399 (1977).Article 

Google Scholar 
Rüber, L., Kottelat, M., Tan, H. H., Ng, P. K. & Britz, R. Evolution of miniaturization and the phylogenetic position of Paedocypris, comprising the world’s smallest vertebrate. BMC Evol. Biol. 7(1), 1–10. https://doi.org/10.1186/1471-2148-7-38 (2007).Article 
CAS 

Google Scholar 
Britz, R., Conway, K. W. & Rüber, L. Miniatures, morphology and molecules: Paedocypris and its phylogenetic position (Teleostei, Cypriniformes). Zool. J. Linn. Soc. 172(3), 556–615. https://doi.org/10.1111/zoj.12184 (2014).Article 

Google Scholar 
Bloom, D. D., Kolmann, M., Foster, K. & Watrous, H. Mode of miniaturisation influences body shape evolution in New World anchovies (Engraulidae). J. Fish Biol. 96(1), 194–201 (2019).Article 

Google Scholar 
Thacker, C. E. Molecular phylogeny of the gobioid fishes (Teleostei: Perciformes: Gobioidei). Mol. Phylogenet. Evol. 26, 354–368. https://doi.org/10.1016/S1055-7903(02)00361-5 (2003).Article 
CAS 

Google Scholar 
Birdsong, R. S., Murdy, E. O. & Pezold, F. L. A study of the vertebral column and median fin osteology in gobioid fishes with comments on gobioid relationships. Bull. Mar. Sci. 42(2), 174–214 (1988).
Google Scholar 
Thacker, C. E. Patterns of divergence in fish species separated by the Isthmus of Panama. BMC Evol. Biol. 17(1), 1–14. https://doi.org/10.1186/s12862-017-0957-4 (2017).Article 

Google Scholar 
Galván-Quesada, S. et al. Molecular phylogeny and biogeography of the amphidromous fish genus Dormitator Gill 1861 (Teleostei: Eleotridae). PLoS One 11(4), e0153538. https://doi.org/10.1371/journal.pone.0153538 (2016).Article 
CAS 

Google Scholar 
Lessios, H. A. The great American schism: Divergence of marine organisms after therise of the central American isthmus. Annu. Rev. Ecol. Evol. Syst. 2008(39), 63–92. https://doi.org/10.1146/annurev.ecolsys.38.091206.095815 (2008).Article 

Google Scholar 
Lovejoy, N. R., Albert, J. S. & Crampton, W. G. Miocene marine incursions and marine/freshwater transitions: Evidence from Neotropical fishes. J. S. Am. Earth Sci. 21, 5–13. https://doi.org/10.1016/j.jsames.2005.07.009 (2006).Article 

Google Scholar 
Cooke, G. M., Chao, N. L. & Beheregaray, L. B. Marine incursions, cryptic species and ecological diversification in Amazonia: The biogeographic history of the croaker genus Plagioscion (Sciaenidae). J. Biogeogr. 39, 724–738. https://doi.org/10.1111/j.1365-2699.2011.02635.x (2012).Article 

Google Scholar 
Bloom, D. D. & Lovejoy, N. R. On the origins of marine-derived freshwater fishes in South America. J. Biogeogr. 44(9), 1927–1938. https://doi.org/10.1111/jbi.12954 (2017).Article 

Google Scholar 
Monsch, K. A. Miocene fish faunas from the northwestern Amazonia basin (Colombia, Peru, Brazil) with evidence of marine incursions. Palaeogeogr. Palaeoclimatol. Palaeoecol. 143, 31–50. https://doi.org/10.1016/S0031-0182(98)00064-9 (1998).Article 

Google Scholar 
Hoorn, C. Marine incursions and the influence of Andean tectonics on the Miocene depositional history of northwestern Amazonia: Results of a palynostratigraphic study. Palaeogeogr. Palaeoclimatol. Palaeoecol. 105, 267–309. https://doi.org/10.1016/0031-0182(93)90087-Y (1993).Article 

Google Scholar 
Hoorn, C., Guerrero, J., Sarmiento, G. A. & Lorente, M. A. Andean tectonics as a cause for changing drainage patterns in Miocene northern South America. Geology 23, 237–240. https://doi.org/10.1130/0091-7613(1995)023%3C0237:ATAACF%3E2.3.CO;2 (1995).Article 
ADS 

Google Scholar 
Gingras, M. K., Rasanen, M. E., Pemberton, S. G. & Romero, L. P. Ichnology and sedimentology reveal depositional characteristics of bay-margin parasequences in the Miocene Amazonian foreland basin. J. Sediment. Res. 72, 871–883. https://doi.org/10.1306/052002720871 (2002).Article 
ADS 

Google Scholar 
Wesselingh, F. P. et al. Lake Pebas: A palaeoecological reconstruction of a Miocene, long-lived lake complex in western Amazonia. Cainoz. Res. 1, 35–81 (2002).
Google Scholar 
Bloom, D. D. & Lovejoy, N. R. Molecular phylogenetics reveals a pattern of biome conservatism in New World anchovies (family Engraulidae). J. Evol. Biol. 25(4), 701–715 (2012).Article 

Google Scholar 
Ward, A. B. & Azizi, E. Convergent evolution of the head retraction escape response in elongate fishes and amphibians. Zoology 107(3), 205–217. https://doi.org/10.1016/j.zool.2004.04.003 (2004).Article 

Google Scholar 
Palumbi, S. R. & Benzie, J. Large mitochondrial DNA differences between morphologically similar penaeid shrimp. Mol. Mar. Biol. Biotechnol. 1, 27–34 (1991).CAS 

Google Scholar 
Chen, W. J., Bonillo, C. & Lecointre, G. Repeatability of clades as criterion of reliability: A case study for molecular phylogeny of Acanthomorpha (Teleostei) with larger number of taxa. Mol. Phylogenet. Evol. 26, 262–288. https://doi.org/10.1016/j.gene.2008.07.016 (2003).Article 
CAS 

Google Scholar 
Chen, W. J., Miya, M., Saitoh, K. & Mayden, R. L. Phylogenetic utility of two existing and four novel nuclear gene loci in reconstructing Tree of Life of ray-finned fishes: The order Cypriniformes (Ostariophysi) as a case study. Gene 423, 125–134. https://doi.org/10.1016/j.gene.2008.07.016 (2008).Article 
CAS 

Google Scholar 
Sanger, F., Nicklen, S. & Coulson, A. R. DNA sequencing with chain-terminating inhibitors. PNAS 74(12), 5463–5467. https://doi.org/10.1073/pnas.74.12.5463 (1977).Article 
ADS 
CAS 

Google Scholar 
Edgar, R. C. MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res. 32(5), 1792–1797. https://doi.org/10.1093/nar/gkh340 (2004).Article 
CAS 

Google Scholar 
Vaidya, G., Lohman, D. J. & Meier, R. SequenceMatrix: Concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics 27, 171–180 (2011).Article 

Google Scholar 
Lanfear, R., Frandsen, P. B., Wright, A. M., Senfeld, T. & Calcott, B. PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Mol. Biol. Evol. https://doi.org/10.1093/molbev/msw260 (2016).Article 

Google Scholar 
Heled, J. & Drummond, A. J. Bayesian inference of population size history from multiple loci. BMC Evol. Biol. 8(1), 1–15. https://doi.org/10.1186/1471-2148-8-289 (2008).Article 
CAS 

Google Scholar 
Bouckaert, R. et al. BEAST 2: A software platform for bayesian evolutionary analysis. PLoS Comput. Biol. 10(4), e1003537. https://doi.org/10.1371/journal.pcbi.1003537 (2014).Article 
CAS 

Google Scholar 
Drummond, A. J., Ho, S. Y., Phillips, M. J. & Rambaut, A. Relaxed phylogenetics and dating with confidence. PLoS Biol. 4(5), e88. https://doi.org/10.1371/journal.pbio.0040088 (2006).Article 
CAS 

Google Scholar 
Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67(5), 901. https://doi.org/10.1093/sysbio/syy032 (2018).Article 
CAS 

Google Scholar 
Drummond, A. J. & Rambaut, A. BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evol. Biol. 7, 214. https://doi.org/10.1186/1471-2148-7-214 (2007).Article 
CAS 

Google Scholar 
Rambaut, A. FigTree, a graphical viewer of phylogenetic trees (Version 1.4.3) (2017).Betancur-R, R. et al. Phylogenetic classification of bony fishes. BMC Evol. Biol. 17(1), 1–40. https://doi.org/10.1186/s12862-017-0958-3 (2017).Article 

Google Scholar 
Jones, G. Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. J. Math. Biol. 74, 447–467 (2017).Article 
MathSciNet 
MATH 

Google Scholar  More

Bovet, D. & Vauclair, J. Picture recognition in animals and humans. Behav. Brain. Res. 109, 143–165 (2000).Article 
CAS 

Google Scholar 
Anonymous. Tinder for Orangutans. Dublin Zoo. https://www.dublinzoo.ie/news/tinder-for-orangutans (2020).Henley, J. “Tinder for Orangutans”: Dutch zoo to let female choose mate on a tablet. The Guardian. https://www.theguardian.com/environment/2017/jan/31/tinder-for-orangutans-dutch-zoo-to-let-female-choose-mate-on-a-tablet (2017).Gierszewski, S. et al. The virtual lover: variable and easily guided 3D fish animations as an innovative tool in mate-choice experiments with sailfin mollies-II validation. Curr. Zool. 6, 65–74 (2017).Article 

Google Scholar 
Dolins, F. L., Klimowicz, C., Kelley, J. & Menzel, C. R. Using virtual reality to investigate comparative spatial cognitive abilities in chimpanzees and humans. Am. J. Primat. 76, 496–513 (2014).Article 

Google Scholar 
Faria, J. J. et al. A novel method for investigating the collective behaviour of fish: Introducing ‘Robofish’. Behav. Ecol. Sociobiol. 64, 1211–1218 (2010).Article 

Google Scholar 
Kozak, E. C. & Uetz, G. W. Male courtship signal modality and female mate preference in the wolf spider Schizocosa ocreata: results of digital multimodal playback studies. Curr. Zool. 65, 705–711 (2019).Article 

Google Scholar 
Loukola, O. J., Perry, C. J., Coscos, L. & Chittka, L. Bumblebees show cognitive flexibility by improving on an observed complex behavior. Science 355, 833–836 (2017).Article 
ADS 
CAS 

Google Scholar 
MacLaren, R. D. Evidence of an emerging female preference for an artificial male trait and the potential for spread via mate choice copying in Poecilia latipinna. Ethology 125, 575–586 (2019).
Google Scholar 
Rönkä, K. et al. Geographic mosaic of selection by avian predators on hindwing warning colour in a polymorphic aposematic moth. Ecol. Lett. 23, 1654–1663 (2020).Article 

Google Scholar 
Rosenthal, G. G., Rand, A. S. & Ryan, M. J. The vocal sac as a visual cue in anuran communication: An experimental analysis using video playback. Anim. Behav. 68, 55–58 (2004).Article 

Google Scholar 
Thurley, K. & Ayaz, A. Virtual reality systems for rodents. Curr. Zool. 63, 109–119 (2017).Article 

Google Scholar 
Ware, E. L., Saunders, D. R. & Troje, N. F. Social interactivity in pigeon courtship behavior. Curr. Zool. 63, 85–95 (2017).Article 

Google Scholar 
Wang, D. et al. The influence of model quality on self-other mate choice copying. Pers. Ind. Diff. 17, 110481 (2021).Article 

Google Scholar 
Gray, J. R., Pawlowski, V. & Willis, M. A. A method for recording behavior and multineuronal CNS activity from tethered insects flying in virtual space. J. Neurosci. Meth. 120, 211–223 (2002).Article 

Google Scholar 
Strauss, R., Schuster, S. & Götz, K. G. Processing of artificial visual feedback in the walking fruit fly Drosophila melanogaster. J. Exp. Biol. 200, 1281–1296 (1997).Article 
CAS 

Google Scholar 
Kemppainen, J. et al. Binocular mirror-symmetric microsaccadic sampling enables Drosophila hyperacute 3D vision. PNAS 119, e2109717119 (2022).Article 
CAS 

Google Scholar 
Bowers, R. I., Place, S. S., Todd, P. M., Penke, L. & Asendorpf, J. B. Generalization in mate-choice copying in humans. Behav. Ecol. 23, 112–124 (2012).Article 

Google Scholar 
Pruett-Jones, S. Independent versus nonindependent mate choice: do females copy each other? Am. Nat. 140, 1000–1006 (1992).Article 
CAS 

Google Scholar 
Dagaeff, A.-C., Pocheville, A., Nöbel, S., Isabel, G. & Danchin, E. Drosophila mate copying correlates with atmospheric pressure in a speed learning situation. Anim. Behav. 121, 163–174 (2016).Article 

Google Scholar 
Mery, F. et al. Public versus personal information for mate copying in an invertebrate. Curr. Biol. 19, 730–734 (2009).Article 
CAS 

Google Scholar 
Danchin, E. et al. Cultural flies: Conformist social learning in fruitflies predicts long-lasting mate-choice traditions. Science 362, 1025–1030 (2018).Article 
ADS 
CAS 

Google Scholar 
Monier, M., Nöbel, S., Isabel, G. & Danchin, E. Effects of a sex ratio gradient on female mate-copying and choosiness in Drosophila melanogaster. Curr. Zool. 64, 251–258 (2018).Article 

Google Scholar 
Monier, M., Nöbel, S., Danchin, E. & Isabel, G. Dopamine and serotonin are both required for mate-copying in Drosophila melanogaster. Front. Behav. Neurosci. 12, 334 (2019).Article 

Google Scholar 
Nöbel, S., Allain, M., Isabel, G. & Danchin, E. Mate copying in Drosophila melanogaster males. Anim. Behav. 141, 9–15 (2018).Article 

Google Scholar 
Nöbel, S., Danchin, E. & Isabel, G. Mate-copying for a costly variant in Drosophila melanogaster females. Behav. Ecol. 29, 1150–1156 (2018).Article 

Google Scholar 
Dukas, R. Natural history of social and sexual behavior in fruit flies. Sci. rep. 10, 1–11 (2020).Article 

Google Scholar 
Chouinard-Thuly, L. et al. Technical and conceptual considerations for using animated stimuli in studies of animal behavior. Curr. Zool. 63, 5–19 (2017).Article 

Google Scholar 
Nöbel, S. et al. Female fruit flies copy the acceptance, but not the rejection, of a mate. Behav. Ecol. 33, 1018–1024 (2022)Article 

Google Scholar 
Bretman, A., Westmancoat, J. D., Gage, M. J. G. & Chapman, T. Males use multiple, redundant cues to detect mating rivals. Curr. Biol. 21, 617–622 (2011).Article 
CAS 

Google Scholar 
Greenspan, R. J. & Ferveur, J. F. Courtship in drosophila. Ann. Rev. Gen. 34, 205 (2000).Article 
CAS 

Google Scholar 
Grillet, M., Dartevelle, L. & Ferveur, J. F. A Drosophila male pheromone affects female sexual receptivity. Proc. Roy. Soc. B. 273, 315–323 (2006).Article 
CAS 

Google Scholar 
Borst, A. Drosophila’s view on insect vision. Curr. Biol. 19, R36–R47 (2009).Article 
CAS 

Google Scholar 
Paulk, A., Millard, S. & van Swinderen, B. Vision in Drosophila: Seeing the world through a model´s eye. Ann. Rev. Entomol. 58, 313–332 (2013).Article 
CAS 

Google Scholar 
Antony, C. & Jallon, J. M. The chemical basis for sex recognition in Drosophila melanogaster. J. Insect. Physiol. 28, 873–880 (1982).Article 
CAS 

Google Scholar 
Keesey, I. W. et al. Adult frass provides a pheromone signature for Drosophila feeding and aggregation. J. Chem. Ecol. 42, 739–747 (2016).Article 
CAS 

Google Scholar 
Talyn, B. C. & Bowse, H. B. The role of courtship song in sexual selection and species recognition by female Drosophila melanogaster. Anim. Behav. 68, 1165–1180 (2004).Article 

Google Scholar 
von Schilcher, F. The function of pulse song and sine song in the courtship of Drosophila melanogaster. Anim. Behav. 24, 622–6251976 (1976).Article 

Google Scholar 
McGregor, P. K. et al. Design of playback experiments: The Thornbridge hall NATO ARW consensus. In Playback and Studies of Animal Communication (ed. McGregor, P.) 1–9 (Plenum Press, New York, 1992).Chapter 

Google Scholar 
Richmond, J. The three Rs. In The UFAW Handbook on the Care and Management of Laboratory and Other Research Animals (eds Hubrecht, R. & Kirkwood, J.) 5–22 (Wiley-Blackwell, Hoboken, 2002).
Google Scholar 
Russell, W. M. S. & Burch, R. L. The Principles of Humane Experimental Technique (Methuen & Co Ltd, 1959).
Google Scholar 
Schlupp, I., Ryan, M. & Waschulewski, M. Female preferences for naturally-occurring novel male traits. Behaviour 136, 519–527 (1999).Article 

Google Scholar 
Witte, K. & Klink, K. No pre-existing bias in sailfin molly females, Poecilia latipinna, for a sword in males. Behaviour 142, 283–303 (2005).Article 

Google Scholar 
Gerlai, R. Animated images in the analysis of zebrafish behavior. Curr. Zool. 63, 35–44 (2017).Article 

Google Scholar 
Ioannou, C. C., Guttal, V. & Couzin, I. D. Predatory fish select for coordinated collective motion in virtual prey. Science 337, 1212–1215 (2012).Article 
ADS 
CAS 

Google Scholar 
Little, A. C., Jones, B. C. & DeBruine, L. M. Preferences for variation in masculinity in real male faces change across the menstrual cycle: Women prefer more masculine faces when they are more fertile. Pers. Ind. Diff. 45, 478–482 (2008).Article 

Google Scholar 
Little, A. C., Jones, B. C. & DeBruine, L. M. Facial attractiveness: Evolutionary based research. Phil. Trans. R. Soc. B. 366, 1638–1659 (2011).Article 

Google Scholar 
Morrison, E. R., Clark, A. P., Tiddeman, B. P. & Penton-Voak, I. S. Manipulating shape cues in dynamic human faces: Sexual dimorphism is preferred in female but not male faces. Ethology 116, 1234–1243 (2010).Article 

Google Scholar 
Kacsoh, B. Z., Bozler, J., Ramaswami, M. & Bosco, G. Social communication of predator-induced changes in Drosophila behavior and germ line physiology. eLife. 4, e07423 (2015).Article 

Google Scholar 
Caruana, N. & Seymour, K. Objects that induce face pareidolia are prioritized by the visual system. Brit. J. Psychol. 113, 496–507 (2022).Article 

Google Scholar 
Agrawal, S., Safarik, S. & Dickinson, M. The relative roles of vision and chemosensation in mate recognition of Drosophila melanogaster. J. Exp. Biol. 217, 2796–2805 (2014).
Google Scholar 
R Development Core Team. R: A Language and Environment for Statistical Computing (Austria, Vienna, 2021).
Google Scholar 
Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).Article 

Google Scholar 
Fox, J. & Weisberg, S. An {R} Companion to Applied Regression 2nd edn. (Sage Publishing, London, 2001).
Google Scholar  More

Richard, E., Said, S., Hamann, J. L. & Gaillard, J. M. Daily, seasonal and annual variations in individual home range overlap of two sympatric spacies of deer. Can. J. Zool. 92, 853–859 (2014).Article 

Google Scholar 
Sorensen, A. A., Stenhouse, G. B., Bourbonnais, M. L. & Nelson, T. A. Effects of habitat quality and anthropogenic disturbance on grizzly bear (Ursus arctos horribilis) home-range fidelity. Can. J. Zool. 93, 857–865 (2015).Article 

Google Scholar 
van Beest, F. M., Rivrud, I. M., Loe, L. E., Milner, J. M. & Mysterud, A. What determines variation in home range size across spatiotemporal scales in a large browsing herbivore?. J. Anim. Ecol. 80, 771–785 (2011).Article 

Google Scholar 
Naidoo, R., Du, P., Weaver, G. S. L. C., Jago, M. & Wegmann, M. Factors affecting intraspecific variation in home range size of a large African herbivore. Landsc. Ecol. 27, 1523–1534 (2012).Article 

Google Scholar 
Bose, S. et al. Implications of fidelity and philopatry for the population structure of female black-tailed deer. Behav. Ecol. 28, 983–990 (2017).Article 

Google Scholar 
Northrup, J. M., Anderson, C. R. Jr. & Wittemyer, G. Environmental dynamics and anthropogenic development alter philopatry and space-use in a North American cervid. Divers. Distrib. 22, 547–557 (2016).Article 

Google Scholar 
Passadore, C., Möller, L., Diaz-aguirre, F. & Parra, G. J. High site fidelity and restricted ranging patterns in southern Australian bottlenose dolphins. Ecol. Evol. 8, 242–256 (2018).Article 

Google Scholar 
Morales, J. M. et al. Building the bridge between animal movement and population dynamics. Philos. Trans. R. Soc. B Biol. Sci. 365, 2289–2301 (2010).Article 

Google Scholar 
Shaw, A. K. Causes and consequences of individual variation in animal movement. Mov. Ecol. 8, 1–12 (2020).Article 

Google Scholar 
Morrison, T. A. et al. Drivers of site fidelity in ungulates. J. Anim. Ecol. 00, 1–12 (2021).
Google Scholar 
Abrahms, B. et al. Emerging perspectives on resource tracking and animal movement ecology. Trends Ecol. Evol. 36, 308–320 (2021).Article 

Google Scholar 
Barraquand, F. & Benhamou, S. Animal movements in heterogeneous landscapes: Identifying profitable places and homogeneous movement bouts. Ecology 89, 3336–3348 (2008).Article 

Google Scholar 
Mueller, T. & Fagan, W. F. Search and navigation in dynamic environments: From individual behaviors to population distributions. Oikos 117, 654–664 (2008).Article 

Google Scholar 
Sawyer, H., Merkle, J. A., Middleton, A. D., Dwinnell, S. P. H. & Monteith, K. L. Migratory plasticity is not ubiquitous among large herbivores. J. Anim. Ecol. 88, 450–460 (2019).
Google Scholar 
Shakeri, Y. N., White, K. S. & Waite, J. N. Staying close to home: Ecological constraints on space use and range fidelity in a mountain ungulate. Ecol. Evol. 11, 11051–11064 (2021).Article 

Google Scholar 
Damuth, J. Home range, home range overlap, and species energy use among herbivorous mammals. Biol. J. Linn. Soc. 15, 185–193 (1981).Article 

Google Scholar 
Lindstedt, S. L., Miller, B. J. & Buskirk, S. W. Home range, time, and body size in mammals. Ecol. Soc. Am. 67, 413–418 (1986).
Google Scholar 
Ofstad, E. G., Herfindal, I., Solberg, E. J. & Sæther, B. E. Home ranges, habitat and body mass: Simple correlates of home range size in ungulates. Proc. R. Soc. B Biol. Sci. 283, 20161234 (2016).Article 

Google Scholar 
Gehr, B. et al. Stay home, stay safe—Site familiarity reduces predation risk in a large herbivore in two contrasting study sites. J. Anim. Ecol. 89, 1329–1339 (2020).Article 

Google Scholar 
Sach, F., Dierenfeld, E. S., Langley-Evans, S. C., Watts, M. J. & Yon, L. African savanna elephants (Loxodonta africana) as an example of a herbivore making movement choices based on nutritional needs. PeerJ 7, 1–27 (2019).Article 

Google Scholar 
Pretorius, Y. et al. Diet selection of African elephant over time shows changing optimization currency. Oikos 121, 2110–2120 (2012).Article 

Google Scholar 
Chamaillé-Jammes, S., Valeix, M. & Fritz, H. Managing heterogeneity in elephant distribution: Interactions between elephant population density and surface-water availability. J. Appl. Ecol. 44, 625–633 (2007).Article 

Google Scholar 
Purdon, A. & van Aarde, R. J. Water provisioning in Kruger National Park alters elephant spatial utilisation patterns. J. Arid Environ. 141, 45–51 (2017).Article 
ADS 

Google Scholar 
Shannon, G., Matthews, W. S., Page, B. R., Parker, G. E. & Smith, R. J. The affects of artificial water availability on large herbivore ranging patterns in savanna habitats: A new approach based on modelling elephant path distributions. Divers. Distrib. 15, 776–783 (2009).Article 

Google Scholar 
Kos, M. et al. Seasonal diet changes in elephant and impala in mopane woodland. Eur. J. Wildl. Res. 58, 279–287 (2012).Article 

Google Scholar 
Shannon, G., Mackey, R. L. & Slotow, R. Diet selection and seasonal dietary switch of a large sexually dimorphic herbivore. Acta Oecologica 46, 48–55 (2013).Article 
ADS 

Google Scholar 
Loarie, S. R., van Aarde, R. J. & Pimm, S. L. Elephant seasonal vegetation preferences across dry and wet savannas. Biol. Conserv. 142, 3099–3107 (2009).Article 

Google Scholar 
Scogings, P. F. et al. Seasonal variations in nutrients and secondary metabolites in semi-arid savannas depend on year and species. J. Arid Environ. 114, 54–61 (2015).Article 
ADS 

Google Scholar 
Birkett, P. J., Vanak, A. T., Muggeo, V. M. R., Ferreira, S. M. & Slotow, R. Animal perception of seasonal thresholds: Changes in elephant movement in relation to rainfall patterns. PLoS ONE 7, 1–8 (2012).Article 

Google Scholar 
Cushman, S. A., Chase, M. & Griffin, C. Elephants in space and time. Oikos 109, 331–341 (2005).Article 

Google Scholar 
Bohrer, G., Beck, P. S., Ngene, S. M., Skidmore, A. K. & Douglas-Hamilton, I. Elephant movement closely tracks precipitation-driven vegetation dynamics in a Kenyan forest-savanna landscape. Mov. Ecol. 2, 1–12 (2014).Article 

Google Scholar 
Purdon, A., Mole, M. A., Chase, M. J. & van Aarde, R. J. Partial migration in savanna elephant populations distributed across southern Africa. Sci. Rep. 8, 1–11 (2018).Article 
CAS 

Google Scholar 
Shannon, G., Page, B. R., Duffy, K. J. & Slotow, R. The ranging behaviour of a large sexually dimorphic herbivore in response to seasonal and annual environmental variation. Austral Ecol. 35, 731–742 (2010).Article 

Google Scholar 
Tsalyuk, M., Kilian, W., Reineking, B. & Getz, W. M. Temporal variation in resource selection of African elephants follows long-term variability in resource availability. Ecol. Monogr. 89, 1–19 (2019).Article 

Google Scholar 
Thaker, M., Prins, H. H. T., Slotow, R., Vanak, A. T. & Gupte, P. R. Fine-scale tracking of ambient temperature and movement reveals shuttling behavior of elephants to water. Front. Ecol. Evol. 7, 1–12 (2019).Article 

Google Scholar 
Govender, N., Trollope, W. S. W. & Van Wilgen, B. W. The effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in South Africa. J. Appl. Ecol. 43, 748–758 (2006).Article 

Google Scholar 
MacFadyen, S., Hui, C., Verburg, P. H. & Van Teeffelen, A. J. A. Spatiotemporal distribution dynamics of elephants in response to density, rainfall, rivers and fire in Kruger National Park, South Africa. Divers. Distrib. 25, 880–894 (2019).Article 

Google Scholar 
Edwards, M. A., Nagy, J. A. & Derocher, A. E. Low site fidelity and home range drift in a wide-ranging, large Arctic omnivore. Anim. Behav. 77, 23–28 (2009).Article 

Google Scholar 
Switzer, P. Site fidelity in predictable and unpredictable habitats. Evol. Ecol. 7, 533–555 (1993).Article 

Google Scholar 
Kranstauber, B., Kays, R., Lapoint, S. D., Wikelski, M. & Safi, K. A dynamic Brownian bridge movement model to estimate utilization distributions for heterogeneous animal movement. J. Anim. Ecol. 81, 738–746 (2012).Article 

Google Scholar 
Kranstauber, B., Smolla, M. & Safi, K. Similarity in spatial utilization distributions measured by the earth mover’s distance. Methods Ecol. Evol. 8, 155–160 (2017).Article 

Google Scholar 
Wartmann, F., Juarez, C. & Fernandez-duque, E. Size, site fidelity, and overlap of home ranges and core areas in the socially monogamous owl monkey (Aotus azarae) of Northern Argentina. Int. J. Primatol. 35, 919–939 (2014).Article 

Google Scholar 
Pringle, R. M. Elephants as agents of habitat creation for small vertebrates at the patch scale. Ecology 89, 26–33 (2008).Article 

Google Scholar 
Valeix, M. et al. Elephant-induced structural changes in the vegetation and habitat selection by large herbivores in an African savanna. Biol. Conserv. 144, 902–912 (2011).Article 

Google Scholar 
Coverdale, T. C. et al. Elephants in the understory: opposing direct and indirect effects of consumption and ecosystem engineering by megaherbivores. Ecology 97, 3219–3230 (2016).Article 

Google Scholar 
Gertenbach, W. Rainfall patterns in the Kruger National Park. Koedoe 23, 35–43 (1980).Article 

Google Scholar 
Venter, F. J., Scholes, R. J. & Eckhardt, H. C. The abiotic template and its associated vegetation pattern. In The Kruger Experience (eds du Toit, J. T. et al.) 83–129 (Island Press, 2003).
Google Scholar 
Young, K. D., Ferreira, S. M. & van Aarde, R. J. The influence of increasing population size and vegetation productivity on elephant distribution in the Kruger National Park. Austral Ecol. 34, 329–342 (2009).Article 

Google Scholar 
Ferreira, S. M., Greaver, C. & Simms, C. Elephant population growth in Kruger National Park, South Africa, under a landscape management approach. Koedoe 59, 1–6 (2017).Article 

Google Scholar 
Brownrigg, R. Package ‘Maps’: Draw Geographical Maps (2022).Kranstauber, B. & Smolla, M. Move: Visualizing and analyzing animal track data. R package version 2.1.0 (2013).R Core Team. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. URL https://www.R-project.org/ (2017).Horne, J. S., Garton, E. O., Krone, S. M. & Lewis, J. S. Analyzing animal movement using Brownian bridges. Ecology 88, 2354–2363 (2007).Article 

Google Scholar 
Wato, Y. A. et al. Movement patterns of African elephants (Loxodonta africana) in a semi-arid savanna suggest that they have information on the location of dispersed water sources. Front. Ecol. Evol. 6, 1–8 (2018).Article 

Google Scholar 
Polansky, L., Kilian, W. & Wittemyer, G. Elucidating the significance of spatial memory on movement decisions by African savannah elephants using state-space models. Proc. R. Soc. B Biol. Sci. 282, 1–7 (2015).
Google Scholar 
Archibald, S. & Scholes, R. J. Leaf green-up in a semi-arid African savanna–separating tree and grass responses to environmental cues. J. Veg. Sci. 18, 583–594 (2007).
Google Scholar 
Majozi, N. P. et al. Analysing surface energy balance closure and partitioning over a semi-arid savanna FLUXNET site in Skukuza, Kruger National Park, South Africa. Hydrol. Earth Syst. Sci. 21, 3401–3415 (2017).Article 
ADS 
CAS 

Google Scholar 
Dodge, S. et al. The environmental-data automated track annotation (Env-DATA) system: Linking animal tracks with environmental data. Mov. Ecol. 1, 1–14 (2013).Article 

Google Scholar 
Didan, K. MOD13Q1 MODIS/terra vegetation indices 16-day L3 global 250m SIN Grid V006. NASA EOSDIS Land Process. DAAC https://doi.org/10.5067/MODIS/MOD13Q1.006 (2015).Redfern, J. V., Grant, C. C., Gaylard, A. & Getz, W. M. Surface water availability and the management of herbivore distributions in an African savanna ecosystem. J. Arid Environ. 63, 406–424 (2005).Article 
ADS 

Google Scholar 
Young, K. D., Ferreira, S. M. & van Aarde, R. J. Elephant spatial use in wet and dry savannas of southern Africa. J. Zool. 278, 189–205 (2009).Article 

Google Scholar 
Goldenberg, S. Z., Douglas-Hamilton, I. & Wittemyer, G. Inter-generational change in African elephant range use is associated with poaching risk, primary productivity and adult mortality. Proc. R. Soc. B Biol. Sci. 285, 1–8 (2018).
Google Scholar 
Woolley, L.-A. et al. Population and individual elephant response to a catastrophic fire in Pilanesberg National Park. PLoS ONE 3, 1–10 (2008).Article 

Google Scholar 
Eby, S. L., Anderson, T. M., Mayemba, E. P. & Ritchie, M. E. The effect of fire on habitat selection of mammalian herbivores: The role of body size and vegetation characteristics. J. Anim. Ecol. 83, 1196–1205 (2014).Article 

Google Scholar 
Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9, 378–400 (2017).Article 

Google Scholar 
Burnham, K. P. & Anderson, D. R. Model Selection and Multimodal Inference: A Practical Information-Theoretic Approach (Springer, 2002).MATH 

Google Scholar 
Mazerolle, M. J. AICcmodavg: Model Selection and Multimodel Inference Based on (Q)AIC(c) (2020).van Moorter, B. et al. Memory keeps you at home: A mechanistic model for home range emergence. Oikos 118, 641–652 (2009).Article 

Google Scholar 
Guldemond, R. A. R., Purdon, A. & van Aarde, R. J. A systematic review of elephant impact across Africa. PLoS ONE 12, 1–12 (2017).Article 

Google Scholar 
Abraham, J. O., Goldberg, E. R., Botha, J. & Staver, A. C. Heterogeneity in African savanna elephant distributions and their impacts on trees in Kruger National Park, South Africa. Ecol. Evol. 11, 5624–5634 (2021).Article 

Google Scholar 
Wall, J., Douglas-Hamilton, I. & Vollrath, F. Elephants avoid costly mountaineering. Curr. Biol. 16, 527–529 (2006).Article 

Google Scholar 
Presotto, A., Fayrer-Hosken, R., Curry, C. & Madden, M. Spatial mapping shows that some African elephants use cognitive maps to navigate the core but not the periphery of their home ranges. Anim. Cogn. 22, 251–263 (2019).Article 

Google Scholar 
Landman, M., Schoeman, D. S., Hall-Martin, A. J. & Kerley, G. I. H. Understanding long-term variations in an elephant piosphere effect to manage impacts. PLoS ONE 7, 1–11 (2012).Article 

Google Scholar 
Fahrig, L. et al. Functional landscape heterogeneity and animal biodiversity in agricultural landscapes. Ecol. Lett. 14, 101–112 (2011).Article 

Google Scholar 
Hamm, M. & Drossel, B. Habitat heterogeneity hypothesis and edge effects in model metacommunities. J. Theor. Biol. 426, 40–48 (2017).Article 
ADS 

Google Scholar 
Katayama, N. et al. Landscape heterogeneity-biodiversity relationship: Effect of range size. PLoS ONE 9, 1–8 (2014).Article 

Google Scholar 
Tews, J. et al. Animal species diversity driven by habitat heterogeneity/diversity: The importance of keystone structures. J. Biogeogr. 31, 79–92 (2004).Article 

Google Scholar 
O’Connor, T. G., Goodman, P. S. & Clegg, B. A functional hypothesis of the threat of local extirpation of woody plant species by elephant in Africa. Biol. Conserv. 136, 329–345 (2007).Article 

Google Scholar 
Codron, J. et al. Elephant (Loxodonta africana) diets in Kruger National Park, South Africa: Spatial and landscape differences. J. Mammal. 87, 27–34 (2006).Article 

Google Scholar 
Mduma, S. A. R., Sinclair, A. R. E. & Hilborn, R. Food regulates the Serengeti wildebeest: A 40-year record. J. Anim. Ecol. 68, 1101–1122 (1999).Article 

Google Scholar 
Ogutu, J. O. & Owen-Smith, N. ENSO, rainfall and temperature influences on extreme population declines among African savanna ungulates. Ecol. Lett. 6, 412–419 (2003).Article 

Google Scholar 
Codron, J. et al. Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces. Oecologia 165, 89–99 (2011).Article 
ADS 

Google Scholar 
Woolley, L.-A., Millspaugh, J. J., Woods, R. J., Page, B. R. & Slotow, R. Intraspecific strategic responses of African elephants to temporal variation in forage quality. J. Wildl. Manag. 73, 827–835 (2009).Article 

Google Scholar 
Dube, K. & Nhamo, G. Evidence and impact of climate change on South African national parks. Potential implications for tourism in the Kruger National Park. Environ. Dev. 33, 1–11 (2020).Article 

Google Scholar 
Tshipa, A. et al. Partial migration links local surface-water management to large-scale elephant conservation in the world’s largest transfrontier conservation area. Biol. Conserv. 215, 46–50 (2017).Article 

Google Scholar 
Nathan, R. et al. Big-data approaches lead to an increased understanding of the ecology of animal movement. Science (80-.) 375, 1–12 (2022).Article 

Google Scholar 
Kays, R., Crofoot, M. C., Jetz, W. & Wikelski, M. Terrestrial animal tracking as an eye on life and planet. Science (80-.) 348, 1222–1232 (2015).Article 
CAS 

Google Scholar 
Mpakairi, K. S., Ndaimani, H., Tagwireyi, P., Zvidzai, M. & Madiri, T. H. Futuristic climate change scenario predicts a shrinking habitat for the African elephant (Loxodonta africana): Evidence from Hwange National Park, Zimbabwe. Eur. J. Wildl. Res. 66, 1–10 (2020).Article 

Google Scholar 
Staver, A. C., Wigley-Coetsee, C. & Botha, J. Grazer movements exacerbate grass declines during drought in an African savanna. J. Ecol. 107, 1482–1491 (2019).Article 

Google Scholar 
Asner, G. P., Vaughn, N., Smit, I. P. J. & Levick, S. Ecosystem-scale effects of megafauna in African savannas. Ecography (Cop.) 39, 240–252 (2016).Article 

Google Scholar 
Shannon, G. et al. Relative impacts of elephant and fire on large trees in a savanna ecosystem. Ecosystems 14, 1372–1381 (2011).Article 

Google Scholar 
Mole, M. A., DÁraujo, S. R., van Aarde, R. J., Mitchell, D. & Fuller, A. Coping with heat: Behavioural and physiological responses of savanna elephants in their natural habitat. Conserv. Physiol. 4, 1–11 (2016).Article 

Google Scholar 
Ncongwane, K. P., Botai, J. O., Sivakumar, V., Botai, C. M. & Adeola, A. M. Characteristics and long-term trends of heat stress for South Africa. Sustainability 13, 1–20 (2021).Article 

Google Scholar 
Lagendijk, G., Mackey, R. L., Page, B. R. & Slotow, R. The effects of herbivory by a mega- and mesoherbivore on tree recruitment in sand forest, South Africa. PLoS ONE 6, 1–9 (2011).Article 

Google Scholar 
Wells, H. B. M. et al. Experimental evidence that effects of megaherbivores on mesoherbivore space use are influenced by species’ traits. J. Anim. Ecol. 90, 2510–2522 (2021).Article 

Google Scholar 
Thaker, M. et al. Minimizing predation risk in a landscape of multiple predators: Effects on the spatial distribution of African ungulates. Ecology 92, 398–407 (2011).Article 

Google Scholar 
Fležar, U. et al. Simulated elephant-induced habitat changes can create dynamic landscapes of fear. Biol. Conserv. 237, 267–279 (2019).Article 

Google Scholar 
Brennan, A. et al. Characterizing multispecies connectivity across a transfrontier conservation landscape. J. Appl. Ecol. 57, 1700–1710 (2020).Article 

Google Scholar 
Roever, C. L., van Aarde, R. J. & Leggett, K. Functional connectivity within conservation networks: Delineating corridors for African elephants. Biol. Conserv. 157, 128–135 (2013).Article 

Google Scholar 
Green, S. E., Davidson, Z., Kaaria, T. & Doncaster, C. P. Do wildlife corridors link or extend habitat? Insights from elephant use of a Kenyan wildlife corridor. Afr. J. Ecol. 56, 860–871 (2018).Article 

Google Scholar  More

FAO. Global forest resources assessment. www.fao.org/publications (2015).Finlayson, M. et al. A Report of the Millennium Ecosystem Assessment. (The Cropper Foundation, 2005).Lal, R., & Lorenz, K. In Recarbonization of the Biosphere: Ecosystems and the Global Carbon Cycle (eds Lal, R., Lorenz, K., Hüttl, R. F., Schneider, B. U. & von Braun, J.) Ch. 9 (Springer, 2012).Gilliam, F. S. Forest ecosystems of temperate climatic regions: from ancient use to climate change. N. Phytologist 212, 871–887 (2016).Article 

Google Scholar 
de Gouvenain, R. C. & Silander, J. A. Temperate forests in Reference Module in Life Sciences (Elsevier, 2017).Keith, S. A., Newton, A. C., Morecroft, M. D., Bealey, C. E. & Bullock, J. M. Taxonomic homogenization of woodland plant communities over 70 years. Proc. R. Soc. B: Biol. Sci. 276, 3539–3544 (2009).Article 

Google Scholar 
Rackham, O. Ancient woodlands: modern threats. N. Phytologist 180, 571–586 (2008).Article 

Google Scholar 
Bernhardt-Römermann, M. et al. Drivers of temporal changes in temperate forest plant diversity vary across spatial scales. Glob. Chang. Biol. 21, 3726–3737 (2015).Article 
ADS 

Google Scholar 
Waller, D. M. & Alverson, W. S. The white-tailed deer: a keystone herbivore. Wildl. Soc. Bull. 25, 217–226 (1997).
Google Scholar 
Ramirez, J. I. Uncovering the different scales in deer–forest interactions. Ecol. Evol. 11, 5017–5024 (2021).Article 

Google Scholar 
Rooney, T. P., Wiegmann, S. M., Rogers, D. A. & Waller, D. M. Biotic impoverishment and homogenization in unfragmented forest understory communities. Conserv. Biol. 18, 787–798 (2004).Stockton, S. A., Allombert, S., Gaston, A. J. & Martin, J. L. A natural experiment on the effects of high deer densities on the native flora of coastal temperate rain forests. Biol. Conserv 126, 118–128 (2005).Article 

Google Scholar 
Hegland, S. J., Lilleeng, M. S. & Moe, S. R. Old-growth forest floor richness increases with red deer herbivory intensity. Ecol. Manag. 310, 267–274 (2013).Article 

Google Scholar 
Simončič, T., Bončina, A., Jarni, K. & Klopčič, M. Assessment of the long-term impact of deer on understory vegetation in mixed temperate forests. J. Veg. Sci. 30, 108–120 (2019).Article 

Google Scholar 
Vild, O. et al. The paradox of long-term ungulate impact: increase of plant species richness in a temperate forest. Appl. Veg. Sci. 20, 282–292 (2017).Article 

Google Scholar 
Russell, F. L., Zippin, D. B. & Fowler, N. L. Effects of white-tailed deer (Odocoileus virginianus) on plants, plant populations and communities: a review. Am. Midl. Nat. 146, 1–26 (2001).Article 

Google Scholar 
Öllerer, K. et al. Beyond the obvious impact of domestic livestock grazing on temperate forest vegetation–A global review. Biol. Conserv. 237, 209–219 (2019).Article 

Google Scholar 
Borer, E. T. et al. Nutrients cause grassland biomass to outpace herbivory. Nat. Commun. 11, 1–8 (2020).Article 
ADS 

Google Scholar 
Kaarlejärvi, E., Eskelinen, A. & Olofsson, J. Herbivores rescue diversity in warming tundra by modulating trait-dependent species losses and gains. Nat. Commun. 8, 1–8 (2017).
Google Scholar 
Simkin, S. M. et al. Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States. Proc. Natl Acad. Sci. USA 113, 4086–4091 (2016).Article 
ADS 
CAS 

Google Scholar 
Bobbink, R. et al. Global assessment of nitrogen deposition effects on terrestrial plant diversity: A synthesis. Ecol. Appl. 20, 30–59 (2010).Article 
CAS 

Google Scholar 
Reinecke, J., Klemm, G. & Heinken, T. Vegetation change and homogenization of species composition in temperate nutrient deficient Scots pine forests after 45 yr. J. Veg. Sci. 25, 113–121 (2014).Article 

Google Scholar 
Speed, J. D. M., Austrheim, G., Kolstad, A. L. & Solberg, E. J. Long-term changes in northern large-herbivore communities reveal differential rewilding rates in space and time. PLoS ONE 14, e0217166 (2019).Article 
CAS 

Google Scholar 
Valente, A. M., Acevedo, P., Figueiredo, A. M., Fonseca, C. & Torres, R. T. Overabundant wild ungulate populations in Europe: management with consideration of socio-ecological consequences. Mamm. Rev. 50, 353–366 (2020).Article 

Google Scholar 
Linnell, J. D. C. et al. The challenges and opportunities of coexisting with wild ungulates in the human-dominated landscapes of Europe’s Anthropocene. Biol. Conserv. 244, 108500 (2020).Waller, D. M. The Herbaceous Layer in Forests of Eastern North America (ed. Gilliam, F.) Ch. 16 (Oxford Univ. Press, 2014).Kerley, G. I. H., Kowalczyk, R. & Cromsigt, J. P. G. M. Conservation implications of the refugee species concept and the European bison: king of the forest or refugee in a marginal habitat? Ecography 35, 519–529 (2011).Svenning, J. C. A review of natural vegetation openness in north-western Europe. Biol. Conserv 104, 133–148 (2002).Article 

Google Scholar 
Sandom, C. J., Ejrnaes, R., Hansen, M. D. D. & Svenning, J. C. High herbivore density associated with vegetation diversity in interglacial ecosystems. Proc. Natl Acad. Sci. USA 111, 4162–4167 (2014).Article 
ADS 
CAS 

Google Scholar 
Ramirez, J. I., Jansen, P. A., den Ouden, J., Goudzwaard, L. & Poorter, L. Long-term effects of wild ungulates on the structure, composition and succession of temperate forests. Ecol. Manag. 432, 478–488 (2019).Article 

Google Scholar 
Ramirez, J. I., Jansen, P. A. & Poorter, L. Effects of wild ungulates on the regeneration, structure and functioning of temperate forests: A semi-quantitative review. Ecol. Manag. 424, 406–419 (2018).Article 

Google Scholar 
Albert, A. et al. Seed dispersal by ungulates as an ecological filter: a trait-based meta-analysis. Oikos 124, 1109–1120 (2015).Article 

Google Scholar 
McNaughton, S. J. Grazing lawns: on domesticated and wild grazers. Am. Nat. 128, 937–939 (1986).Article 

Google Scholar 
Cromsigt, J. P. G. M. & Kuijper, D. P. J. Revisiting the browsing lawn concept: evolutionary Interactions or pruning herbivores? Perspect. Plant Ecol. 13, 207–215 (2011).Article 

Google Scholar 
Ramirez, J. I. et al. Temperate forests respond in a non-linear way to a population gradient of wild deer. Forestry 94, 502–511 (2021).Article 

Google Scholar 
Boulanger, V. et al. Ungulates increase forest plant species richness to the benefit of non‐forest specialists. Glob. Chang. Biol. 24, e485–e495 (2018).Article 

Google Scholar 
Kirby, K. J. The impact of deer on the ground flora of British broadleaved woodland. Forestry 74, 219–229 (2001).Article 

Google Scholar 
Royo, A. A., Collins, R., Adams, M. B., Kirschbaum, C. & Carson, W. P. Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity. Ecology 91, 93–105 (2010).Happonen, K. et al. Trait-based responses to land use and canopy dynamics modify long-term diversity changes in forest understories. Glob. Ecol. Biogeogr. 30, 1863–1875 (2021).Article 

Google Scholar 
Peñuelas, J. & Sardans, J. The global nitrogen-phosphorus imbalance. Science 375, 266–267 (2022).Article 
ADS 

Google Scholar 
Staude, I. R. et al. Replacements of small- by large-ranged species scale up to diversity loss in Europe’s temperate forest biome. Nat. Ecol. Evol. 4, 802–808 (2020).Article 

Google Scholar 
Newbold, T. et al. Widespread winners and narrow-ranged losers: Land use homogenizes biodiversity in local assemblages worldwide. PLoS Biol. 16, e2006841 (2018).Article 

Google Scholar 
Verheyen, K. et al. Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests. Br. Ecol. Soc. J. Ecol. 100, 352–365 (2012).
Google Scholar 
Gilliam, F. S. Response of the herbaceous layer of forest ecosystems to excess nitrogen deposition. J. Ecol. 94, 1176–1191 (2006).Article 
CAS 

Google Scholar 
de Schrijver, A. et al. Cumulative nitrogen input drives species loss in terrestrial ecosystems. Glob. Ecol. Biogeogr. 652, 803–816 (2011).Article 

Google Scholar 
de Frenne, P. et al. Light accelerates plant responses to warming. Nat. Plants 1, 15110 (2015).Article 

Google Scholar 
Baeten, L. et al. Herb layer changes (1954-2000) related to the conversion of coppice-with-standards forest and soil acidification. Appl. Veg. Sci. 12, 187–197 (2009).Article 

Google Scholar 
Becker, T., Spanka, J., Schröder, L. & Leuschner, C. Forty years of vegetation change in former coppice-with-standards woodlands as a result of management change and N deposition. Appl. Veg. Sci. 20, 304–313 (2017).Article 

Google Scholar 
van Calster, H. et al. Diverging effects of overstorey conversion scenarios on the understorey vegetation in a former coppice-with-standards forest. Ecol. Manag. 256, 519–528 (2008).Article 

Google Scholar 
Luyssaert, S. et al. The European carbon balance. Part 3: forests. Glob. Chang. Biol. 16, 1429–1450 (2010).Article 
ADS 

Google Scholar 
Kirby, K. J. et al. Five decades of ground flora changes in a temperate forest: the good, the bad and the ambiguous in biodiversity terms. Ecol. Manag. 505, 119896 (2022).Article 

Google Scholar 
Hautier, Y., Niklaus, P. A. & Hector, A. Competition for light causes plant biodiversity loss after eutrophication. Science 324, 636–638 (2009).Article 
ADS 
CAS 

Google Scholar 
Kowalczyk, R., Kamiński, T. & Borowik, T. Do large herbivores maintain open habitats in temperate forests? For. Ecol. Manag. 494, 119310 (2021).Dormann, C. F. et al. Plant species richness increases with light availability, but not variability, in temperate forests understorey. BMC Ecol. 20, 1–9 (2020).Article 

Google Scholar 
Dirnböck, T. et al. Forest floor vegetation response to nitrogen deposition in Europe. Glob. Chang. Biol. 20, 429–440 (2014).Article 
ADS 

Google Scholar 
Perring, M. P. et al. Understanding context dependency in the response of forest understorey plant communities to nitrogen deposition. Environ. Pollut. 242, 1787–1799 (2018).Article 
CAS 

Google Scholar 
Anderson, T. M. et al. Herbivory and eutrophication mediate grassland plant nutrient responses across a global climatic gradient. Ecology 99, 822–831 (2018).Article 

Google Scholar 
Gough, L. & Grace, J. B. Herbivore effects on plant species density at varying productivity levels. Ecology 79, 1586–1594 (1998).Article 

Google Scholar 
Eskelinen, A., Harpole, W. S., Jessen, M.-T., Virtanen, R. & Hautier, Y. Light competition drives herbivore and nutrient effects on plant diversity. Nature 611, 301–305 (2022).Knight, T. M., Dunn, J. L., Smith, L. A., Davis, J. A. & Kalisz, S. Deer facilitate invasive plant success in a Pennsylvania forest understory. Nat. Areas 29, 110–116 (2009).Article 

Google Scholar 
Beguin, J., Pothier, D. & Côté, S. D. Deer browsing and soil disturbance induce cascading effects on plant communities: a multilevel path analysis. Ecol. Appl. 21, 439–451 (2011).Gilliam, F. S. et al. Twenty-five-year response of the herbaceous layer of a temperate hardwood forest to elevated nitrogen deposition. Ecosphere 7, e01250 (2016).Article 

Google Scholar 
de Frenne, P. et al. Microclimate moderates plant responses to macroclimate warming. Proc. Natl Acad. Sci. USA 110, 18561–18565 (2013).Article 
ADS 

Google Scholar 
Hedwall, P. O. et al. Half a century of multiple anthropogenic stressors has altered northern forest understory plant communities. Ecol. Appl. 29, e01874 (2019).Perring, M. P. et al. Global environmental change effects on plant community composition trajectories depend upon management legacies. Glob. Chang. Biol. 24, 1722–1740 (2018).Article 
ADS 

Google Scholar 
Boulanger, V. et al. Decreasing deer browsing pressure influenced understory vegetation dynamics over 30 years. Ann. Sci. 72, 367–378 (2015).Article 

Google Scholar 
Bernes, C. et al. Manipulating ungulate herbivory in temperate and boreal forests: effects on vegetation and invertebrates. A systematic review. Environ. Evid. 7, 1–32 (2018).Article 

Google Scholar 
Reimoser, F. Steering the impacts of ungulates on temperate forests. J. Nat. Conserv. 10, 243–252 (2003).Article 

Google Scholar 
Vavra, M., Parks, C. G. & Wisdom, M. J. Biodiversity, exotic plant species, and herbivory: the good, the bad, and the ungulate. Ecol. Manag. 246, 66–72 (2007).Article 

Google Scholar 
Depauw, L. et al. Light availability and land-use history drive biodiversity and functional changes in forest herb layer communities. J. Ecol. 108, 1411–1425 (2020).Article 
CAS 

Google Scholar 
Chevaux, L. et al. Effects of stand structure and ungulates on understory vegetation in managed and unmanaged forests. Ecol. Appl. 32, e01874 (2022).Gordon, I. J. Browsing and grazing ruminants: are they different beasts? Ecol. Manag. 181, 13–21 (2003).Article 

Google Scholar 
Brasseur, B. et al. What deep‐soil profiles can teach us on deep‐time pH dynamics after land use change? Land Degrad. Dev. 29, 2951–2961 (2018).Article 

Google Scholar 
Schmitz, A. et al. Responses of forest ecosystems in Europe to decreasing nitrogen deposition. Environ. Pollut. 244, 980–994 (2019).Article 
CAS 

Google Scholar 
Dirnböck, T. et al. Currently legislated decreases in nitrogen deposition will yield only limited plant species recovery in European forests. Environ. Res. Lett. 13, 125010 (2018).Article 

Google Scholar 
Peterken, G. F. Natural Woodland: Ecology and Conservation in Northern Temperate Regions (Cambridge Univ. Press, 1996).Chamberlain, S. A. & Boettiger, C. R Python, and Ruby clients for GBIF species occurrence data. preprint. PeerJ Preprints 5, e3304v1 (2017).Chamberlain, S. A. & Szöcs, E. taxize: taxonomic search and retrieval in R. F1000Res 2, 191 (2013).Article 

Google Scholar 
Hédl, R., Kopecký, M. & Komárek, J. Half a century of succession in a temperate oakwood: from species-rich community to mesic forest. Divers Distrib. 16, 267–276 (2010).Article 

Google Scholar 
Giménez-Anaya, A., Herrero, J., Rosell, C., Couto, S. & García-Serrano, A. Food habits of wild boars (Sus scrofa) in a Mediterranean coastal wetland. Wetlands 28, 197–203 (2008).Article 

Google Scholar 
Barrios-Garcia, M. N. & Ballari, S. A. Impact of wild boar (Sus scrofa) in its introduced and native range: a review. Biol. Invasions 14, 2283–2300 (2012).Article 

Google Scholar 
Andersen, R. et al. An overview of the progress and challenges of peatland restoration in Western Europe. Restor. Ecol. 25, 271–282 (2017).Article 

Google Scholar 
Faurby, S. et al. PHYLACINE 1.2: the phylogenetic atlas of mammal macroecology. Ecology 99, 2626 (2018).Article 

Google Scholar 
van den Berg, L. J. L. et al. Evidence for differential effects of reduced and oxidised nitrogen deposition on vegetation independent of nitrogen load. Environ. Pollut. 208, 890–897 (2016).Article 

Google Scholar 
McNaughton, S. J., Oesterheld, M., Frank, D. A. & Williams, K. J. Ecosystem-level patterns of primary productivity and herbivory in terrestrial habitats. Nature 341, 142–144 (1989).Article 
ADS 
CAS 

Google Scholar 
Koerner, S. E. et al. Change in dominance determines herbivore effects on plant biodiversity. Nat. Ecol. Evol. 2, 1925–1932 (2018).Article 

Google Scholar 
Fréjaville, T. & Garzón, M. B. The EuMedClim database: yearly climate data (1901-2014) of 1 km resolution grids for Europe and the Mediterranean Basin. Front. Ecol. Evol. 6, 1–5 (2018).Article 

Google Scholar 
Al‐Yaari, A. et al. Asymmetric responses of ecosystem productivity to rainfall anomalies vary inversely with mean annual rainfall over the conterminous United States. Glob. Chang. Biol. 26, 6959–6973 (2020).Article 
ADS 

Google Scholar 
Szabó, P. & Hédl, R. Advancing the integration of history and ecology for conservation. Conserv. Biol. 25, 680–687 (2011).Article 

Google Scholar 
Hedges, L. V., Gurevitch, J. & Curtis, P. S. The meta-analysis of response ratios in experimental ecology. Spec. Feature Ecol. 80, 1150–1156 (1999).
Google Scholar 
Hillebrand, H. et al. Biodiversity change is uncoupled from species richness trends: consequences for conservation and monitoring. J. Appl. Ecol. 55, 169–184 (2018).Article 

Google Scholar 
Holz, H., Segar, J., Valdez, J. & Staude, I. R. Assessing extinction risk across the geographic ranges of plant species in Europe. Plants People Planet 4, 303–311 (2022).Article 

Google Scholar 
Staude, I. R. et al. Directional turnover towards larger‐ranged plants over time and across habitats. Ecol. Lett. 25, 466–482 (2021).Article 

Google Scholar 
Ellenberg, H., Weber, H. E., Düll, R., Wirth, V. & Werner, W. Zeigerwerte von Pflanzen in Mitteleuropa (Verlag Wrich Goltze, 2001).Chytrý, M., Tichý, L., Dřevojan, P., Sádlo, J. & Zelený, D. Ellenbergtype indicator values for the Czech flora. Preslia 90, 83–103 (2018).Bürkner, P.-C. brms: an R package for Bayesian multilevel models using Stan. J. Stat. Softw. 80, 1–28 (2017).Article 

Google Scholar 
Brooks, S. P. & Gelman, A. General methods for monitoring convergence of iterative simulations. J. Comput. Graph. Stat. 7, 434–455 (1998).MathSciNet 

Google Scholar 
Dushoff, J., Kain, M. P. & Bolker, B. M. I can see clearly now: reinterpreting statistical significance. Methods Ecol. Evol. 10, 756–759 (2019).Article 

Google Scholar 
Bradshaw, L. & Waller, D. M. Impacts of white-tailed deer on regional patterns of forest tree recruitment. Ecol. Manag. 375, 1–11 (2016).Article 

Google Scholar 
McGarvey, J. C., Bourg, N. A., Thompson, J. R., McShea, W. J. & Shen, X. Effects of twenty years of deer exclusion on woody vegetation at three life-history stages in a mid-atlantic temperate deciduous forest. Northeast. Nat. 20, 451–468 (2013).Nuttle, T., Ristau, T. E. & Royo, A. A. Long-term biological legacies of herbivore density in a landscape-scale experiment: forest understoreys reflect past deer density treatments for at least 20 years. J. Ecol. 102, 221–228 (2013). More

Correction to: Communications Biology https://doi.org/10.1038/s42003-020-1022-1, published online 11 June 2020.In the original version of the Perspective, a unit conversion error affected calculations for cereal rye, triticale, barley, and oats. Further, berseem clover yield estimates were mistranscribed from the original source. These mistakes led to errors in Supplementary Data 1, Figure 2 and in the presentation of the data in the text.Supplementary Data 1 has now been replaced with a file containing the correct numbers.Figure 2 has been corrected:Original figure 2New figure 2The Abstract stated: “In this Perspective, we estimate land use requirements to supply the United States maize production area with cover crop seed, finding that across 18 cover crops, on average 3.8% (median 2.0%) of current production area would be required, with the popular cover crops rye and hairy vetch requiring as much as 4.5% and 11.9%, respectively”.The text should read: “In this Perspective, we estimate land use requirements to supply the United States maize production area with cover crop seed, finding that across 18 cover crops, on average 2.4% (median 2.1%) of current production area would be required, with the popular cover crops rye and hairy vetch requiring as much as 4.8% and 11.9%, respectively”.In the 1st paragraph of the right hand column on page 2, the text said: “(…), we find that the land requirements for production of cover crop seed would be on average 1.4 million hectares (median 746,000 ha), which is equivalent to 3.8% (median 2.0%) of the U.S. maize farmland. Rye (Secale cereale L.) – a midrange seed yielding cover crop and one of the most commonly used in the corn belt, would require as much as 1,661,000 hectares (4.5% of maize farmland), (…)”The text should read: “(…) we find that the land requirements for production of cover crop seed would be on average 892,526 hectares (median 774,417 ha), which is equivalent to 2.4% (median 2.1%) of the U.S. maize farmland. Rye (Secale cereale L.) – a midrange seed yielding cover crop and one of the most commonly used in the corn belt, would require as much as 1,779,770 hectares (4.8% of maize farmland), (…)”On page 3, second paragraph the text said: “Cover cropping the entire U.S. maize area would require the equivalent of as much as 18% (rye) to 49% (hairy vetch) (…)”The text should read: “Cover cropping the entire U.S. maize area would require the equivalent of as much as 19% (rye) to 49% (hairy vetch) (…)”This errors have now been corrected in the Perspective Article. More