Philippa Kaur

Zeebe, R. E. & Wolf-Gladrow, D. CO2 in Seawater: Equilibrium, Kinetics, Isotopes (Elsevier, 2001).Ridgwell, A. & Zeebe, R. The role of the global carbonate cycle in the regulation and evolution of the Earth system. Earth Planet. Sci. Lett. 234, 299–315 (2005).Article 

Google Scholar 
Moore, C. M. et al. Processes and patterns of oceanic nutrient limitation. Nat. Geosci. 6, 701–710 (2013).Article 

Google Scholar 
Klausmeier, C. A., Litchman, E., Daufresne, T. & Levin, S. A. Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton. Nature 429, 171–174 (2004).Article 

Google Scholar 
Krumhardt, K. M., Lovenduski, N. S., Iglesias-Rodriguez, M. D. & Kleypas, J. A. Coccolithophore growth and calcification in a changing ocean. Prog. Oceanogr. 159, 276–295 (2017).Article 

Google Scholar 
Zondervan, I. The effects of light, macronutrients, trace metals and CO2 on the production of calcium carbonate and organic carbon in coccolithophores—a review. Deep Sea Res. Part 2 54, 521–537 (2007).Article 

Google Scholar 
Gibbs, S. J., Sheward, R. M., Bown, P. R., Poulton, A. J. & Alvarez, S. A. Warm plankton soup and red herrings: calcareous nannoplankton cellular communities and the Palaeocene–Eocene Thermal Maximum. Phil. Trans. R. Soc. A 376, 20170075 (2018).Article 

Google Scholar 
Aloisi, G. Covariation of metabolic rates and cell size in coccolithophores. Biogeosciences 12, 6215–6284 (2015).Article 

Google Scholar 
Boudreau, B. P., Middelburg, J. J. & Luo, Y. The role of calcification in carbonate compensation. Nat. Geosci. 11, 894–900 (2018).Article 

Google Scholar 
Suchéras-Marx, B. & Henderiks, J. Downsizing the pelagic carbonate factory: impacts of calcareous nannoplankton evolution on carbonate burial over the past 17 million years. Glob. Planet. Change 123, 97–109 (2014).Article 

Google Scholar 
Beaufort, L. et al. Sensitivity of coccolithophores to carbonate chemistry and ocean acidification. Nature 476, 80–83 (2011).Article 

Google Scholar 
McClelland, H. L. O., Bruggeman, J., Hermoso, M. & Rickaby, R. E. M. The origin of carbon isotope vital effects in coccolith calcite. Nat. Commun. 8, 14511 (2017).Article 

Google Scholar 
Bolton, C. T. et al. Decrease in coccolithophore calcification and CO2 since the middle Miocene. Nat. Commun. 7, 10284 (2016).Article 

Google Scholar 
McClelland, H. L. O. et al. Calcification response of a key phytoplankton family to millennial-scale environmental change. Sci. Rep. 6, 34263 (2016).Article 

Google Scholar 
Duchamp-Alphonse, S. et al. Enhanced ocean–atmosphere carbon partitioning via the carbonate counter pump during the last deglacial. Nat. Commun. 9, 2396 (2018).Article 

Google Scholar 
Si, W. & Rosenthal, Y. Reduced continental weathering and marine calcification linked to late Neogene decline in atmospheric CO2. Nat. Geosci. 12, 833–838 (2019).Article 

Google Scholar 
Meier, K. J. S., Berger, C. & Kinkel, H. Increasing coccolith calcification during CO2 rise of the penultimate deglaciation (Termination II). Mar. Micropaleontol. 112, 1–12 (2014).Article 

Google Scholar 
Su, X., Liu, C. & Beaufort, L. Late Quaternary coccolith weight variations in the northern South China Sea and their environmental controls. Mar. Micropaleontol. 154, 101798 (2020).Article 

Google Scholar 
Berger, C., Meier, K. J. S., Kinkel, H. & Baumann, K.-H. Changes in calcification of coccoliths under stable atmospheric CO2. Biogeosciences 11, 929–944 (2014).Article 

Google Scholar 
Zachos, J., Dickens, G. R. & Zeebe, R. E. An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature 451, 279–283 (2008).Article 

Google Scholar 
Foster, G. L., Royer, D. L. & Lunt, D. J. Future climate forcing potentially without precedent in the last 420 million years. Nat. Commun. 8, 14845 (2017).Article 

Google Scholar 
Anagnostou, E. et al. Proxy evidence for state-dependence of climate sensitivity in the Eocene greenhouse. Nat. Commun. 11, 4436 (2020).Article 

Google Scholar 
Holtz, L.-M., Wolf-Gladrow, D. & Thoms, S. Stable carbon isotope signals in particulate organic and inorganic carbon of coccolithophores—a numerical model study for Emiliania huxleyi. J. Theor. Biol. 420, 117–127 (2017).Article 

Google Scholar 
Hermoso, M., Horner, T. J., Minoletti, F. & Rickaby, R. E. M. Constraints on the vital effect in coccolithophore and dinoflagellate calcite by oxygen isotopic modification of seawater. Geochim. Cosmochim. Acta 141, 612–627 (2014).Article 

Google Scholar 
Hermoso, M., Chan, I. Z. X., McClelland, H. L. O., Heureux, A. M. C. & Rickaby, R. E. M. Vanishing coccolith vital effects with alleviated carbon limitation. Biogeosciences 13, 301–312 (2016).Article 

Google Scholar 
Rickaby, R. E. M., Henderiks, J. & Young, J. N. Perturbing phytoplankton: response and isotopic fractionation with changing carbonate chemistry in two coccolithophore species. Clim. Past 6, 771–785 (2010).Article 

Google Scholar 
Ziveri, P. et al. Stable isotope ‘vital effects’ in coccolith calcite. Earth Planet. Sci. Lett. 210, 137–149 (2003).Article 

Google Scholar 
Bolton, C. T. & Stoll, H. M. Late Miocene threshold response of marine algae to carbon dioxide limitation. Nature 500, 558–562 (2013).Article 

Google Scholar 
Henderiks, J. Coccolithophore size rules—reconstructing ancient cell geometry and cellular calcite quota from fossil coccoliths. Mar. Micropaleontol. 67, 143–154 (2008).Article 

Google Scholar 
Sheward, R. M., Poulton, A. J., Gibbs, S. J., Daniels, C. J. & Bown, P. R. Physiology regulates the relationship between coccosphere geometry and growth phase in coccolithophores. Biogeosciences 14, 1493–1509 (2017).Article 

Google Scholar 
Gibbs, S. J. et al. Species-specific growth response of coccolithophores to Palaeocene–Eocene environmental change. Nat. Geosci. 6, 218–222 (2013).Article 

Google Scholar 
Herrmann, S. & Thierstein, H. R. Cenozoic coccolith size changes—evolutionary and/or ecological controls? Palaeogeogr. Palaeoclimatol. Palaeoecol. 333–334, 92–106 (2012).Article 

Google Scholar 
Young, J. R. & Ziveri, P. Calculation of coccolith volume and its use in calibration of carbonate flux estimates. Deep-Sea Research II 22, 1679–1700 (2000).Article 

Google Scholar 
Daniels, C. J., Sheward, R. M. & Poulton, A. J. Biogeochemical implications of comparative growth rates of Emiliania huxleyi and Coccolithus species. Biogeosciences 11, 6915–6925 (2014).Article 

Google Scholar 
Westerhold, T. et al. An astronomically dated record of Earth’s climate and its predictability over the last 66 million years. Science 369, 1383–1387 (2020).Article 

Google Scholar 
Pälike, H. et al. A Cenozoic record of the equatorial Pacific carbonate compensation depth. Nature 488, 609–614 (2012).Article 

Google Scholar 
Misra, S. & Froelich, P. N. Lithium isotope history of cenozoic seawater: changes in silicate weathering and reverse weathering. Science 335, 818–823 (2012).Article 

Google Scholar 
Ravizza, G. E. & Zachos, J. C. in Treatise on Geochemistry Vol. 6 (ed. Elderfield, H.) 551–581 (Elsevier, 2003).McArthur, J. M., Howarth, R. J. & Bailey, T. R. Strontium isotope stratigraphy: LOWESS version 3: best fit to the marine Sr‐isotope curve for 0–509 Ma and accompanying look‐up table for deriving numerical age. J. Geol. 109, 155–170 (2001).Article 

Google Scholar 
Pegram, W. J., Krishnaswami, S., Ravizza, G. E. & Turekian, K. K. The record of sea water 1870s/1860s variation through the Cenozoic. Earth Planet. Sci. Lett. 113, 569–576 (1992).Article 

Google Scholar 
Shipboard Scientific Party, 2004. Leg 208 summary. In Zachos, J. C., Kroon, D. & Blum, P., et al., Proceedings of the Ocean Drilling Program, Initial Reports, 208, 1–112: College Station, TX (Ocean Drilling Program) (2004).Brummer, G. J. A. & van Eijden, A. J. M. “Blue-ocean” paleoproductivity estimates from pelagic carbonate mass accumulation rates. Mar. Micropaleontol. 19, 99–117 (1992).Article 

Google Scholar 
Gafar, N. A., Eyre, B. D. & Schulz, K. G. A conceptual model for projecting coccolithophorid growth, calcification and photosynthetic carbon fixation rates in response to global ocean change. Front. Mar. Sci. 4, 433 (2018).Article 

Google Scholar 
Gafar, N. A. & Schulz, K. G. A three-dimensional niche comparison of Emiliania huxleyi and Gephyrocapsa oceanica: reconciling observations with projections. Biogeosciences 15, 3541–3560 (2018).Article 

Google Scholar 
Gafar, N. A., Eyre, B. D. & Schulz, K. G. A comparison of species specific sensitivities to changing light and carbonate chemistry in calcifying marine phytoplankton. Sci. Rep. 9, 2486 (2019).Article 

Google Scholar 
Zhang, Y. G. et al. Refining the alkenone–pCO2 method I: lessons from the Quaternary glacial cycles. Geochim. Cosmochim. Acta 260, 177–191 (2019).Article 

Google Scholar 
Freeman, K. H. & Pagani, M. in A History of Atmospheric CO2 and Its Effects on Plants, Animals, and Ecosystems Vol. 177 (eds Baldwin, I. T. et al.) 35–61 (Springer-Verlag, 2005).Pagani, M. The alkenone–CO2 proxy and ancient atmospheric carbon dioxide. Phil. Trans. R. Soc. A 360, 609–632 (2002).Article 

Google Scholar 
Beerling, D. J. & Royer, D. L. Convergent Cenozoic CO2 history. Nat. Geosci. 4, 418–420 (2011).Article 

Google Scholar 
Henehan, M. J. et al. Revisiting the Middle Eocene Climatic Optimum ‘Carbon Cycle Conundrum’ with new estimates of atmospheric pCO2 from boron isotopes. Paleoceanogr. Paleoclimatol. https://doi.org/10.1029/2019PA003713 (2020).Zachos, J., Pagani, M., Sloan, L. C., Thomas, E. & Billups, K. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686–693 (2001).Article 

Google Scholar 
Stap, L., Sluijs, A., Thomas, E. & Lourens, L. Patterns and magnitude of deep sea carbonate dissolution during Eocene Thermal Maximum 2 and H2, Walvis Ridge, southeastern Atlantic Ocean, Paleoceanography 24, PA1211, https://agupubs.onlinelibrary.wiley.com/doi/abs/10.1029/2008PA001655 (2009).Sluijs, A. et al. Warm and wet conditions in the Arctic region during Eocene Thermal Maximum 2. Nat. Geosci. 2, 777–780 (2009).Article 

Google Scholar 
Stap, L. et al. High-resolution deep-sea carbon and oxygen isotope records of Eocene Thermal Maximum 2 and H2. Geology 38, 607–610 (2010).Article 

Google Scholar 
Bohaty, S. M. & Zachos, J. C. Significant Southern Ocean warming event in the late middle Eocene. Geology 31, 1017 (2003).Article 

Google Scholar 
van der Ploeg, R. et al. Middle Eocene greenhouse warming facilitated by diminished weathering feedback. Nat. Commun. 9, 2877 (2018).Article 

Google Scholar 
Bach, L. T., Riebesell, U., Gutowska, M. A., Federwisch, L. & Schulz, K. G. A unifying concept of coccolithophore sensitivity to changing carbonate chemistry embedded in an ecological framework. Prog. Oceanogr. 135, 125–138 (2015).Article 

Google Scholar 
Monteiro, F. M. et al. Why marine phytoplankton calcify. Sci. Adv. 2, e1501822–e1501822 (2016).Article 

Google Scholar 
Shipboard Scientific Party, 2004. Site 1263. In Zachos, J. C., Kroon, D., Blum, P., et al., Proceedings of the Ocean Drilling Program, Initial Reports, 208, 1–87 College Station, TX (Ocean Drilling Program) (2004).Bice, K. L., Sloan, L. C. & Barron, E. J. in Warm Climates in Earth History (eds Huber, B. T., Macleod, K. G., & Wing, S. L.) 79–129 (Cambridge Univ. Press, 2000).Handoh, I. C., Bigg, G. R. & Jones, E. J. W. Evolution of upwelling in the Atlantic Ocean basin. Palaeogeogr. Palaeoclimatol. Palaeoecol. 202, 31–58 (2003).Article 

Google Scholar 
Minoletti, F., Hermoso, M. & Gressier, V. Separation of sedimentary micron-sized particles for palaeoceanography and calcareous nannoplankton biogeochemistry. Nat. Protoc. 4, 14–24 (2009).Article 

Google Scholar 
Zhang, H., Stoll, H., Bolton, C., Jin, X. & Liu, C. A refinement of coccolith separation methods: Measuring the sinking characters of coccoliths. Biogeosciences Discussions (2018): 1–30 https://doi.org/10.5194/bg-2018-82 (2020).Hermoso, M. et al. Towards the use of the coccolith vital effects in palaeoceanography: a field investigation during the middle Miocene in the SW Pacific Ocean. Deep Sea Res. Part 1 160, 103262 (2020).Article 

Google Scholar 
Lauretano, V., Hilgen, F. J., Zachos, J. C. & Lourens, L. J. Astronomically tuned age model for the early Eocene carbon isotope events: a new high-resolution δ13Cbenthic record of ODP site 1263 between ~49 and ~54 Ma. Newsl. Stratigr. 49, 383–400 (2016).Article 

Google Scholar 
Westerhold, T., Röhl, U., Frederichs, T., Bohaty, S. M. & Zachos, J. C. Astronomical calibration of the geological timescale: closing the middle Eocene gap. Clim. Past 11, 1181–1195 (2015).Article 

Google Scholar 
Westerhold, T. et al. Astronomical Calibration of the Ypresian Time Scale: Implications for Seafloor Spreading Rates and the Chaotic Behaviour of the Solar System? Preprint at Clim. Past Discuss. https://doi.org/10.5194/cp-2017-15 (2017).Gatuso, J. P., Epitalon, J. M., Lavigne, H. & Orr, J. seacarb: Seawater Carbonate Chemistry (2021); https://CRAN.R-project.org/package=seacarb More

Ngo, L. T., Okogun, J. I. & Folk, W. R. 21st century natural product research and drug development and traditional medicines. Nat. Prod. Rep. 30, 584–592 (2013).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Li, S. P., Wu, D. T., Lv, G. P. & Zhao, J. Carbohydrates analysis in herbal glycomics. Trac-Trends Anal. Chem. 52, 155–169 (2013).Article 
CAS 

Google Scholar 
Lei, H. B. et al. A comprehensive quality evaluation of Fuzi and its processed product through integration of UPLC-QTOF/MS combined MS/MS-based mass spectral molecular networking with multivariate statistical analysis and HPLC-MS/MS. J. Ethnopharmacol. 266, 113455 (2021).CAS 
PubMed 
Article 

Google Scholar 
Zhao, Z. Z. et al. A unique issue in the standardization of Chinese materia medica: Processing. Planta Med. 76, 1975–1986 (2010).CAS 
PubMed 
Article 

Google Scholar 
Zotz, G. The systematic distribution of vascular epiphytes-a critical update. Bot. J. Lin. Soc. 171, 453–481 (2013).Article 

Google Scholar 
Li, X. et al. Genetic diversity analysis and conservation of the endangered Chinese endemic herb Dendrobium officinale Kimura et Migo (Orchidaceae) based on AFLP. Genetica 133, 159–166 (2008).CAS 
PubMed 
Article 

Google Scholar 
Xu, M. et al. Transcriptome sequencing and development of novel genic SSR markers for Dendrobium officinale. Mol. Breed 37, 1–7 (2017).Article 
CAS 

Google Scholar 
Ng, T. B. et al. Review of research on Dendrobium, a prized folk medicine. Appl. Microbiol. Biotechnol. 93(5), 1795–1803 (2012).CAS 
PubMed 
Article 

Google Scholar 
Zhao, Y. J., Han, B. X., Peng, H. S. & Peng, D. Y. Research on evolution and transition of quality evaluation of Shihu. China J. Chin. Mater. Med. 41, 1348–1353 (2016).
Google Scholar 
Kim, S. N. et al. Simultaneous quantification of 14 ginsenosides in Panax ginseng CA Meyer (Korean red ginseng) by HPLC-ELSD and its application to quality control. J. Pharm. Biomed. Anal. 45, 164–170 (2007).CAS 
PubMed 
Article 

Google Scholar 
Csupor, D. et al. Qualitative and quantitative analysis of aconitine-type and lipo-alkaloids of Aconitum carmichaelii roots. J. Chromatogr. A 1216, 2079–2086 (2009).CAS 
PubMed 
Article 

Google Scholar 
Wan, J. Y. et al. Integrated evaluation of malonyl ginsenosides, amino acids and polysaccharides in fresh and processed ginseng. J. Pharm. Biomed. Anal. 107, 89–97 (2015).CAS 
PubMed 
Article 

Google Scholar 
Li, S. L. et al. Chemical profiling of Radix Paeoniae evaluated by ultra-performance liquid chromatography/photo-diode-array/quadrupole time-of-flight mass spectrometry. J. Pharm. Biomed. Anal. 49, 253–266 (2009).PubMed 
Article 
CAS 

Google Scholar 
Sumner, L. W., Lei, Z., Nikolau, B. J. & Saito, K. Modern plant metabolomics: Advanced natural product gene discoveries, improved technologies, and future prospects. Nat. Prod. Rep. 32, 212–229 (2015).CAS 
PubMed 
Article 

Google Scholar 
Hu, C. & Xu, G. Metabolomics and traditional Chinese medicine. TrAC-Trend Anal. Chem. 61, 207–214 (2014).CAS 
Article 

Google Scholar 
Zhang, A. et al. Metabolomics: Towards understanding traditional Chinese medicine. Planta Med. 76, 2026–2035 (2010).CAS 
PubMed 
Article 

Google Scholar 
Sun, H. et al. Metabolomics study on Fuzi and its processed products using ultra-performance liquid-chromatography/electrospray-ionization synapt high-definition mass spectrometry coupled with pattern recognition analysis. Analyst 137, 170–185 (2012).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Wang, X. et al. Metabolomics study on the toxicity of aconite root and its processed products using ultraperformance liquid-chromatography/electrospray-ionization synapt high-definition mass spectrometry coupled with pattern recognition approach and ingenuity pathways analysis. J. Proteome Res. 11, 1284–1301 (2012).CAS 
PubMed 
Article 

Google Scholar 
Geng, L. et al. Discrimination of raw and vinegar-processed Genkwa Flos using metabolomics coupled with multivariate data analysis: A discrimination study with metabolomics coupled with PCA. Fitoterapia 84, 286–294 (2013).CAS 
PubMed 
Article 

Google Scholar 
Chen, W. H. et al. Traditional uses, phytochemistry, pharmacology, and quality control of Dendrobium offificinale Kimura et. Migo. Front Pharmacol. 12, 726528 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Wang, Y., Tong, Y., Adejobi, O. I., Wang, Y. H. & Liu, A. Z. Research advances in multi-omics on the traditional Chinese Herb Dendrobium offificinale. Front. Phant Sci. 12, 808288 (2022).
Google Scholar 
Yue, H., Zeng, H. & Ding, K. A review of isolation methods, structure features and bioactivities of polysaccharides from Dendrobium species. Chin. J. Nat. Med. 18, 1–27 (2020).CAS 
PubMed 

Google Scholar 
Fahy, E. et al. Update of the LIPID MAPS comprehensive classification system for lipids1. J. Lipid Res. 50, S9–S14 (2009).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Surmacz, L. & Swiezewska, E. Polyisoprenoids–secondary metabolites or physiologically important superlipids?. Biochemi Biophys. Res. Commun. 407, 627–632 (2011).CAS 
PubMed 
Article 

Google Scholar 
Kidd, P. M. Vitamins D and K as pleiotropic nutrients: Clinical importance to the skeletal and cardiovascular systems and preliminary evidence for synergy. Altern. Med. Rev. 15, 199–222 (2010).PubMed 

Google Scholar 
Jiang, Q. Natural forms of vitamin E: Metabolism, antioxidant, and anti-inflammatory activities and their role in disease prevention and therapy. Free Radic. Biol. Medi. 72, 76–90 (2014).CAS 
Article 

Google Scholar 
Bayat, P., Farshchi, M., Yousefian, M., Mahmoudi, M. & Yazdian-Robati, R. Flavonoids, the compounds with anti-inflammatory and immunomodulatory properties, as promising tools in multiple sclerosis (MS) therapy: A systematic review of preclinical evidence. Int. Immunopharmacol. 95, 107562 (2021).CAS 
PubMed 
Article 

Google Scholar 
Heim, K. E., Tagliaferro, A. R. & Bobilya, D. J. Flavonoid antioxidants: Chemistry, metabolism and structure-activity relationships. J. Nutrit. Biochem. 13, 572–584 (2002).CAS 
Article 

Google Scholar 
Aquila, S., Giner, R. M., Recio, M. C., Spegazzini, E. D. & Ríos, J. L. Anti-inflammatory activity of flavonoids from Cayaponia tayuya roots. J. Ethnopharmacol. 121, 333–337 (2009).CAS 
PubMed 
Article 

Google Scholar 
Sharma, H., Kumar, P., Deshmukh, R. R., Bishayee, A. & Kumar, S. Pentacyclic triterpenes: New tools to fight metabolic syndrome. Phytomedicine 50, 166–177 (2018).CAS 
PubMed 
Article 

Google Scholar 
Hodon, J., Borkova, L., Pokorny, J., Kazakova, A. & Urban, M. Design and synthesis of pentacyclic triterpene conjugates and their use in medicinal research. Eur. J. Medi. Chem. 182, 111653 (2019).CAS 
Article 

Google Scholar 
Huang, J. et al. Pentacyclic triterpene carboxylic acids derivatives integrated piperazine-amino acid complexes for α-glucosidase inhibition in vitro. Bioorg. Chem. 115, 105212 (2021).CAS 
PubMed 
Article 

Google Scholar 
Toldrá, F. & Flores, M. The role of muscle proteases and lipases in flavor development during the processing of dry-cured ham. Crit. Rev. Food Sci. Nutr. 38, 331–352 (1998).PubMed 
Article 

Google Scholar 
Imm, J., Zhang, G., Chan, L. Y., Nitteranon, V. & Parkin, K. L. [6]-Dehydroshogaol, a minor component in ginger rhizome, exhibits quinone reductase inducing and anti-inflammatory activities that rival those of curcumin. Food Res. Int. 43, 2208–2213 (2010).CAS 
Article 

Google Scholar 
Surh, Y. J. & Na, H. K. NF-kB and Nrf2 as prime molecular targets for chemoprevention and cytoprotection with anti-inflammatory and antioxidant phytochemicals. Genes Nutr. 2, 313–317 (2008).CAS 
PubMed 
Article 

Google Scholar 
Pan, M. H., Lai, C. S., Dushenkov, S. & Ho, C. T. Modulation of inflammatory genes by natural dietary bioactive compounds. J. Agric. Food Chem. 57, 4467–4477 (2009).CAS 
PubMed 
Article 

Google Scholar 
Privitera, R. & Anand, P. Capsaicin 8% patch Qutenza and other current treatments for neuropathic pain in chemotherapy-induced peripheral neuropathy (CIPN). Curr. Opin Support. Palliat. Car 15, 125–131 (2021).Article 

Google Scholar 
Vila, D. L., Nunes, N., Almeida, P., Gomes, J., Rosa, C., & Alvarez-Leite, J. I. Signaling targets related to antiobesity effects of capsaicin: A scoping review. Adv. Nutr. 1–12 (2021).Simpson, D. M., Estanislao, L., Brown, S. J. & Sampson, J. An open-label pilot study of high-concentration capsaicin patch in painful HIV neuropathy. J. Pain Symptom Manag. 35(3), 299–306 (2008).CAS 
Article 

Google Scholar 
Westphal, C., Cermax, J., Cole, R. O., Short, G. F., Perni, R., & Ponduru, S. Formulations and use of TRP channel activators in treatment of nervous system disorders. PCT Int. Appl. WO 2015160843 A1 (2015).Chen, C. L., Mao, C., Zhang, J. T. Application of vanilloid receptor agonist to prepare anti-Alzheimer’s medical products. Faming Zhuanli Shenqing. CN 1736485 A (2006).Andoh, R., Sakurada, S., Kisara, K., Takahashi, M. & Ohsawa, K. Effects of intra-arterially administered capsaicinoids on vocalization in guinea pigs and medial thalamic neuronal activity in cats. Nippon Yakurigaku Zasshi 79, 275–283 (1982).CAS 
PubMed 
Article 

Google Scholar 
Chen, M. et al. Ligustrum robustum (Roxb.) blume extract modulates gut microbiota and prevents metabolic syndrome in high-fat diet-fed mice. J. Ethnopharmacol. 268, 113695 (2021).CAS 
PubMed 
Article 

Google Scholar 
Kasai, T. & Sakamura, S. Acidic α-acylarginine derivatives in apple and pear trees. Phytochemistry 23, 19–22 (1984).CAS 
Article 

Google Scholar 
Kuley, E., Kuscu, M. M., Durmus, M. & Ucar, Y. Inhibitory activity of Co-microencapsulation of cell free supernatant from Lactobacillus plantarum with propolis extracts towards fish spoilage bacteria. LWT-Food. Sci. Technol. 146, 111433 (2021).CAS 
Article 

Google Scholar 
Adna, M. et al. Vincenzo, Phytochemistry, bioactivities, pharmacokinetics and toxicity prediction of Selaginella repanda with its anticancer potential against human lung, breast and colorectal carcinoma cell lines. Molecules 26, 768 (2021).CAS 
Article 

Google Scholar 
Edenharder, R., Keller, G., Platt, K. L. & Unger, K. K. Isolation and characterization of structurally novel antimutagenic flavonoids from spinach (Spinacia oleracea). J. Agric. Food Chem. 49, 2767–2773 (2001).CAS 
PubMed 
Article 

Google Scholar 
Kumar, V. et al. Chemopreventive effects of Melastoma malabathricum L. extract in mammary tumor model via inhibition of oxidative stress and inflammatory cytokines. Biomed. Pharmacother. 137, 111298 (2021).CAS 
PubMed 
Article 

Google Scholar 
Miceli, N. et al. Phytochemical profile and antioxidant activity of the aerial part extracts from matthiola incana subsp. rupestris and subsp. pulchella (Brassicaceae) endemic to sicily. Chem. Biodivers. 18, e2100167 (2021).CAS 
PubMed 
Article 

Google Scholar 
Pateliya, B., Burade, V. & Goswami, S. Enhanced antitumor activity of doxorubicin by naringenin and metformin in breast carcinoma: An experimental study. Naunyn-Schmiedebergs Arch. Pharmacol. 394, 1949–1961 (2021).CAS 
PubMed 
Article 

Google Scholar 
Kuo, P. C. et al. Anti-inflammatory principles from the needles of Pinus taiwanensis Hayata and in silico studies of their potential anti-aging effects. Antioxidants 10, 598 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Waring, R., & Hunter, J. Methods and antibodies for detecting and measuring toxins in feces for diagnosis of equine laminitis and therapeutic applications. PCT Int. Appl. WO 2019186141 A1 (2019).Oliveira, I. et al. New lectins from Mediterranean flora. Activity against HT29 colon cancer cells. Int. J. Mol. Sci. 20, 3059 (2019).CAS 
PubMed Central 
Article 

Google Scholar 
Choi, A., Nam, Y. H., Baek, K. & Chung, E. J. Brevibacillus antibioticus sp. Nov., with a broad range of antibacterial activity, isolated from soil in the Nakdong River. J. Microbiol. 57, 991–996 (2019).CAS 
PubMed 
Article 

Google Scholar 
Adnan, M. et al. Evaluation of anti-nociceptive and anti-inflammatory activities of the methanol extract of Holigarna caustica (Dennst.) Oken leaves. J. Ethnopharmacol. 236, 401–411 (2019).CAS 
PubMed 
Article 

Google Scholar 
He, S. Z. et al. Hydroxysafflor yellow a inhibits staphylococcus aureus-induced mouse endometrial inflammation via TLR2-Mediated NF-kB and MAPK pathway. Inflammation 44, 835–845 (2021).CAS 
PubMed 
Article 

Google Scholar 
Zhou, W. et al. Network pharmacology to explore the anti-inflammatory mechanism of Xuebijing in the treatment of sepsis. Phytomedicine 85, 153543 (2021).CAS 
PubMed 
Article 

Google Scholar 
Zhao, F. et al. Hydroxysafflower yellow A: A systematical review on botanical resources, physicochemical properties, drug delivery system, pharmacokinetics, and pharmacological effects. Front. Pharmacol. 11, 579332 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Hu, L. J. et al. Dratanguticumides G and H, two new glucosides from Dracocephalum tanguticum Maxim relax vessels via NO pathway. Phytochem. Lett. 40, 42–48 (2020).CAS 
Article 

Google Scholar 
Guo, R. et al. A new sesquiterpenoid with cytotoxic and anti-inflammatory activity from the leaves of Datura metel L. Nat. Prod. Res. 35(4), 607–613 (2021).CAS 
PubMed 
Article 

Google Scholar 
Bai, Y. et al. Comparison of phenolic compounds, antioxidant and antidiabetic activities between selected edible beans and their different growth periods leaves. J. Funct. Food. 35, 694–702 (2017).CAS 
Article 

Google Scholar 
Ouyang, X. L., Wei, L. X., Wang, H. S. & Pan, Y. M. Antioxidant activity and phytochemical composition of Osmanthus fragrans’ pulps. S. Afr. J. Bot. 98, 162–166 (2015).CAS 
Article 

Google Scholar 
Li, Y. et al. Antidepressant-like effects of Cistanche tubulosa extract on chronic unpredictable stress rats through restoration of gut microbiota homeostasis. Front. Pharmacol. 9, 967 (2018).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Chen, Z. M., Yin, M. M., Wang, Z., Liu, H. Y., & Wang, M. L. Application of pharmaceutical composition for preparing medicine for treating gastric cancer based on regulation and control of tumor suppressor-related genes. Faming Zhuanli Shenqing. CN 112716935 A (2021).Leyva-Jimenez, F. J. et al. Incorporation of Lippia citriodora microwave extract into total-green biogelatin-phospholipid vesicles to improve its antioxidant activity. Nanomaterials 10, 765 (2020).CAS 
PubMed Central 
Article 

Google Scholar 
Wu, X. et al. Effect of phenolic hydroxyl groups on inhibitory activities of phenylpropanoid glycosides against lipase. J. Funct. Food 38, 510–518 (2017).Article 
CAS 

Google Scholar 
Lin, Y. E. et al. Antidepressant-like effects of water extract of Cordyceps militaris (Linn.) Link by modulation of ROCK2/PTEN/Akt signaling in an unpredictable chronic mild stress-induced animal model. J. Ethnopharmacol. 276, 114194 (2021).CAS 
PubMed 
Article 

Google Scholar 
Haron, M. H. et al. Effect of African potato (Hypoxis hemerocallidea) extract and its Constituents on PXR and CYP450 enzymes. Appl. In Vitro Toxicol. 5, 26–33 (2019).CAS 
Article 

Google Scholar 
Kamzolova, S. V., Samoilenko, V. A., LuninaIgor, J. N. & Morgunov, G. Effects of medium components on isocitric acid production by Yarrowia lipolytica yeast. Fermentation 6, 112 (2020).CAS 
Article 

Google Scholar 
Strzepek-Gomolka, M. et al. Identification of mushroom and murine tyrosinase inhibitors from Achillea biebersteinii Afan. Extract. Mol. 26, 964 (2021).CAS 

Google Scholar 
Lachowicz, S. et al. Impact mineralization of chokeberry and cranberry fruit juices using a new functional additive on the protection of bioactive compounds and antioxidative properties. Molecules 25, 659 (2020).CAS 
PubMed Central 
Article 

Google Scholar 
Badary, O. A., Nagi, M. N., Al-Sawaf, H. A., Al-Harbi, M. & Al-Bekairi, A. M. Effect of L-histidinol on cisplatin nephrotoxicity in the rat. Nephron 77, 435–439 (1997).CAS 
PubMed 
Article 

Google Scholar 
Badary, O. A. L-histidinol attenuates Fanconi syndrome induced by ifosfamide in rats. Exp. Nephrol. 7, 323–327 (1999).CAS 
PubMed 
Article 

Google Scholar 
Al-Gharably, N. M. & Al-Sawaf, H. A. Effects of L-histidinol on the antitumor activity and acute cardiotoxicity of doxorubicin in mice. Pharmacol. Res. 38, 225–230 (1998).PubMed 
Article 

Google Scholar 
Eze, F. N. & Tola, A. J. Protein glycation and oxidation inhibitory activity of Centella asiatica phenolics (CAP) in glucose-mediated bovine serum albumin glycoxidation. Food Chem. 332, 127302 (2020).CAS 
PubMed 
Article 

Google Scholar 
Peng, Y. et al. Metabolomics study of the anti-inflammatory effects of endogenous omega-3 polyunsaturated fatty acids. RSC Adv. 9, 41903–41912 (2019).ADS 
CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Li, H. G. et al. Study on the nutritional characteristics and antioxidant activity of dealcoholized sequentially fermented apple juice with Saccharomyces cerevisiae and Lactobacillus plantarum fermentation. Food Chem. 363, 130351 (2021).CAS 
PubMed 
Article 

Google Scholar 
Liu, Y. Research on jade fungus sport beverage and its anti-exercise fatigue function. Food Res. Dev. 38, 106–110 (2017).CAS 

Google Scholar 
Zeitoun, H., Khan, Z., Banerjee, K., Salameh, D. & Lteif, R. Antityrosinase activity of Combretum micranthum, Euphorbia hirta and Anacardium occidentale plants: Ultrasound assisted extraction optimization and profiling of associated predominant metabolites. Molecules 25, 2684 (2020).CAS 
PubMed Central 
Article 

Google Scholar 
Rajaram, R., Muralisankar, T., Paray, B. A. & Al-Sadoon, M. K. Phytochemical profiling and antioxidant capacity of Kappaphycus alvarezii (Doty) Doty collected from seaweed farming sites of tropical coastal environment. Aquac. Res. 52, 3438–3448 (2021).CAS 
Article 

Google Scholar 
Basholli-Salihu, M. et al. Phytochemical composition, anti-inflammatory activity and cytotoxic effects of essential oils from three Pinus spp. Pharm. Biol. 55, 1553–1560 (2017).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
He, Z.D. et al. Antioxidative glucosides from the fruits of Ligustrum lucidum. Chem. Pharm. Bull. 49, 780–784 (2001).CAS 
Article 

Google Scholar  More

The Ecomobility is a cross-sectoral, global partnership, an environmentally friendly and socially inclusive way of transportation, seeking the promotion integration of cycling, walking, and passenging (Being a passenger in another’s car or a carpool)1, and wheeling (wheelchairs, non-motorized scooters, walking aids, skates, push scooters, trailer, hand carts, shopping carts/ trolleys, carrying aids)2. In this section, the study introduces the main pillars that will build on them to reach the objectives. Passenging, energy efficiency, partnership for sustainable urban development, and streets’ design (with more focus as one of the core prerequisites of the methods) are discussed here. Although this research approaches the clean and green environment from the window of excluding the fossil-fuel motorized means of transportation, if there is still a necessity, passenging owns a pack of advantages over private cars. As a main passenging option, Carpooling has a lot of benefits to the environment, human health, and social lives, e.g., saving money, better for the environment, Convenience, etc., which are briefed below.PassengingBeing a passenger in another’s car or carpooling that increases efficiency. The concept becomes more apparent, listing the benefits of carpooling as follows3:

1.

Saving money: sharing the cost of gas and parking, cutting expenses by nearly 50%; the more occupants in carpooling, the more saving, and is socially economical. Also, there will be a reduction in constructing new roads, maintenance, and air pollution-related health costs.

2.

Better environment: fewer cars on roads mean reduced Greenhouse Gas (GHG) emissions and better air quality.

3.

Better health: helps reduce some health risks for a human being, such as cardiovascular and respiratory diseases, allergies, and neurological effects.

4.

Convenience: provides commuting Convenience, less stress, and added advantage of companionship while commuting.

5.

Better commuting options: works better for people living where transit service may be non-existent or limited.

Energy efficiencyA study was performed in China to reduce GHG; the study claimed that China targeted the peak around 20304. However, it is expected that the peak may come earlier than expected, which is reflected in the CO2 emission (from coal consumption and fossil fuels) to have its peak also before 2030. Another study evaluated the economic impacts of GHG emission reduction on the Brazilian economy reached that different sectoral targets may balance environmental benefits with the possible financial losses incurred by taxation policy or emission permits5.The energy efficiency studies touched key triggers to reduce carbon emissions, In parallel. An analytical study in China (2013) indicated that if greater intensity emission reduction measures were taken, the carbon emissions would reduce by 31.01 million tons by 2015 and 48.81 million tons by 20206. The previous results indicated the average reduction flow per time.Partnership for sustainable urban developmentSaving the environment is cooperative work that requires sharing experiences with other partners worldwide. The cooperation between the Gulf region and China in the last half of the decade is an example of a global partnership. The Gulf region has achieved a success story, economically and socially. Following a smart investment arrangement of natural resources, the region has succeeded in attracting and retaining international experience, therefore overcoming the discrepancy between its vast economy and small population-base lacking the instantly needed technical skills. The region has collected considerable wealth during the process, financially and else7.The primary purpose of Ecomobility, mentioned previously, is enhancing the opportunity for sustainable urban development. Also, it complies with John Elkington’s triple bottom line (TBL), or what is often referred to as the three “P’s”: people, planet, and prosperity8. Likewise, worldwide, large cities are applying guidelines to guarantee that environment, economics, and sociality are at the lead of urban design. The elevation of healthier streets has formed new chances for social and commercial interaction and more comprehensive outcomes9. Overall, it is the transformation of the earlier global Coalitions of Ecomobility that aims at engaging public and private sectors, promoting and advocating Ecomobility at a worldwide level in Industrialized and developing countries.Streets’ designA potential approach to fostering pedestrian Ecomobility is the streets’ design, which is a core focus of this research. Urban spaces are a vital city part, forming the basic structure of public life. Specific urban planning and design criteria make these spaces “quality public spaces”10. In this perspective, determining and evaluating these criteria will transform those public urban spaces into quality spaces. The last research adopted criteria that came to a head in the relevant reviews and were accepted by most researchers. Also, in the current research, the study chose following these criteria since they have been collected and validated by several researchers over decades (around 17 references; Table 1), and cover the possible aspects of successful streets; they will be presented and used in the next section (Methods).Table 1 Respondents’ statistics (N = 257).Full size tableA noteworthy point is the gap between municipal policies and the guidance provided to street design decision-makers. A study across cities in the United States highlighted some issues regarding the mechanism of developing policies related, as they aim to challenge auto-centrical street design standards in favor of “complete streets,” which are safe for users of all abilities. Also, they address the demands of non-motorized street users and sustainable transportation. Moreover, those policies do not lead to the negotiation of tradeoffs among users inside the street right-of-way; They are broad and defer to optimistic safety goals accommodating all user types equally without recognizing the accommodation’s implicit hierarchy11.Also, the transformative potential of experimentations proposed attaining “streets for people” rather than “streets for traffic” remains under-investigated. There is little to no comparative assessment of already present trials and no critical reflection on their explicit added value for systemic change. While street research aims to create basically diverse arrangements of urban mobility, their potential as triggers of a larger systemic change is blurred12.In the age of autonomous vehicles (AVs), a study reached that with the promise of Avs, which will use less street-right of way and uncouple parking from land uses in the cities; it is time to take back streets and make them serve people first, arranging cycling, walking, and transit. This action will take place precisely in the design process for livable streets. Also, it is time to act and make sure that designers and planners can get at the forefront of the rapidly changing technologies of vehicles13.The street design process had a different approach through medical and psychological professionals. According to the Irvine-Minnesota audit, expert observers calculated street users for four streets, which differed in walkability. From 7 am to 7 pm all days, the whole streets had significant quadratic trends of increasing followed by decreasing use. Furthermore, the two most walkable streets showed substantial linear increases in users across the day. Part of a street’s identity is its temporal activity rhythm, and both walkability and rhythms can report to urban design and restoration14. The study, through its test, searched what makes streets and neighborhoods walkable and found that more walkable streets had more users overall and linear increases in use starting from morning to early evening.This paper focuses on Ecomobility in Riyadh city streets that promote the integration of passenging, cycling, wheeling, and walking. A previous study discussed the pedestrian mobility status in the same city, focusing on the transformation to sustainable mobility. That research case study was applied to the PSU community and argued that the current mobility is unsustainable, as it profoundly relies on privately-owned cars run with fossil fuel. It unveiled that a substantial percentage of the survey sample (72%) traveled by car from home to the campus. However, the assessment results showed that the transformation to sustainable mobility is expected soon by launching the new mega projects related to public transportation like Riyadh Metro and busses, which is considered a key indicator of sustainable mobility15.Riyadh city in Saudi Arabia is a part of a hot-weather zone (almost 7 months a year: April–October), as seen in Fig. 116. As a result, studying the pedestrian thermal comfort affected by street design is a core point. Almost similar conditions were present in a case study in the Australian North Melbourne (southeast) at street level for pedestrians (Subtropical Zone), as seen in Fig. 217. That study assessed modeled existing and future scenarios for different street profiles and the consequences of microclimatic parameters and thermal comfort. The target was to assist urban planners in developing policies that can efficiently reduce the exposure to heat stress at the pedestrian level18.Figure 1The climate in Riyadh16.Full size imageFigure 2World climate zones17.Full size imageA comparison study was also applied to two gulf cities that share the hot weather: Dubai and Abu Dhabi. It compared the efficiency of the early suburbs and the newer ones to provide quick and direct access to destinations (connectivity). The study argued that better connectivity is required for usefulness in hot, dry regions. In this regard, it explored how the abandoned system of alleys could be cultivated to enhance connectivity efficiency19.In residential streets, developers’ width is not a choice but somewhat a constraint enforced through planners’ concluded subdivision standards. Residents can generally compromise by choosing smaller yards or homes in a swap for other facilities or a lower price, but they cannot select a smaller street20.Recently, in Riyadh city, a study highlighted that the rising fuel prices would create a positive atmosphere for implementing the complete streets’ concept, defined as streets’ design that can safely accommodate all transport modes for all society segments. It includes, but is not limited to, public transportation, humanizing neighborhoods, and promoting walking as a healthy lifestyle21.Research problemLacking the dependency on the Ecomobility traveling modes (as mentioned in the results) leads to poor health conditions, air pollution, noise, high greenhouse gas (and CO2) emissions, higher expenses, and excessive energy consumption.Research questionsThe study is seeking through its parts to answer the following questions:

1.

What are the reasons behind the lack of using the ecomobility means of transportation by the community?

2.

What are the community responses for the transformation to ecomobility if the reported barriers are removed or minimized? More

Perched among the fronds of the world’s loneliest tree, Viswambharan Sarasan had an important decision to make. Sarasan had worked for years to get access to this palm — the last living member of the species Hyophorbe amaricaulis, which grows in Curepipe Botanic Gardens, Mauritius.He reached up towards a cluster of its walnut-sized, olive-green fruit. Sarasan, a botanist at the Royal Botanic Gardens at Kew, near London, had been through sensitive negotiations for permission to take the fruit, each with one crucial seed inside. He then had to wait for the tree, nicknamed the lonesome palm, to produce them. Nine metres up, 50 fruit dangling within his grasp, he had to decide how many to take: enough to give himself a chance of culturing them back at Kew, while leaving enough for local scientists to work with.“It was the only shot I could get,” he says of his visit in June 2006. “But I didn’t want to take all the seeds and then it turns out badly.”He picked ten fruit. It was not his lucky number.When the plight of trees gets publicity, deforestation is generally the reason, but it is not the only crisis they face. Nearly one-third of trees — more than 17,500 species — are threatened with extinction. This is more than twice the number of threatened mammals, birds, amphibians and reptiles combined1. Mass plantings of trees, paradoxically, often add to the problem by using single species. Now, hundreds of plant conservationists globally are fighting to save the trees speeding towards extinction.“We shouldn’t be giving up on any tree species,” says Paul Smith, head of Botanic Gardens Conservation International (BGCI), a London-based charity that co-leads the campaign to secure the future of the world’s threatened tree species.But time is short, the obstacles are formidable and both climate change and fashions in ecology are moving against them.Peter Bridgewater, a specialist in biodiversity governance at the University of Canberra, Australia, says that finding a natural home for every tree species is impossible because climate change is altering ecosystems so fundamentally. Scientists who think this goal is realistic are “living in their own cloud cuckoo land”, he says.

The ‘lonesome palm’, which lives in the Curepipe Botanic Gardens in Mauritius, is the last surviving member of the species Hyophorbe amaricaulis. Researchers have tried for years to help it reproduce, without success.Credit: Vincent Florens

Neglected trees Inextricably linked with the problem of climate change, and equally as damaging, is the disappearance of species from Earth. The rate of extinction is at historic levels and accelerating, with around one million animals and plants under threat.The plight of trees can get lost among the tales of endangered mammals or birds. To get trees more visibility, in 2016 the BGCI, working with the International Union for Conservation of Nature (IUCN), organized the largest conservation assessment in the IUCN’s history: the Global Tree Assessment. Hundreds of plant conservationists searched rainforests, mountains and strife-torn regions, sometimes with no more than a crinkly herbarium specimen or the testimony of a long-dead explorer to guide them.In a 2021 report, they announced that they had found 58,497 tree species, of which 17,510 were threatened2. Since then, almost 2,800 of those have been labelled as critically endangered. Some 142 species are thought to be extinct in the wild (see ‘Trees under threat’). This year, a separate group of modellers estimated that a further 9,000 tree species are undiscovered3.

Source: Ref. 2

It is not just the number of trees, but also their diversity that matters. A single species can be the foundation of an entire ecological network, and its disappearance could cause a cascade of extinctions that might lead to an ecosystem collapse.Strong, diverse ecosystems are also better at sequestering carbon, says Jean-Christophe Vié, director-general of the Franklinia Foundation, a private organization in Geneva, Switzerland, that funds tree conservation and supports the Global Tree Assessment. No tree species should be viewed as dispensable, says Vié, because it would set a precedent for every developer, farmer or logger to justify removing any threatened tree.But tree conservation has become lost in international biodiversity targets — partly because trees get subsumed into general plant-conservation goals, and because plants are generally less showy than birds and animals. Trees need to be assessed for ecologists to champion them, says Malin Rivers, head of conservation prioritization at the BGCI.“If you look at mammals, birds, reptiles, they have data to bring to the table when there is a policy discussion,” she says. “Taxonomy gives the species a name; conservation assessment gives it a voice.”Protect and propagateArmed with the Global Tree Assessment’s catalogue of threatened species, conservationists have begun prioritizing species and taxonomic groups. The best approach, says Smith, is to protect vulnerable trees in their natural habitats. If that’s not possible, researchers try growing them from seed in a laboratory, greenhouse or botanic garden.The Global Tree Assessment revealed that nearly two-thirds of threatened trees are found in areas that are already protected, and stressed that one important task is to strengthen or expand these havens.That might mean controlling grazing, implementing a national logging ban for a particular species or establishing plots on which the tree can be cultivated for fruit or flowers without harming the larger population. On the eastern Caribbean island of Dominica, for instance, where harvesting resin for incense was killing lansan trees (Protium attenuatum), a tweak to the tapping method has halted the damage.Sometimes, however, so few trees are left that protecting an area isn’t enough.

In its forest habitat in Tanzania, Karomia gigas is threatened by a seed-killing fungus.Credit: Kirsty Shaw/BGCI

In Tanzania, seed-biology specialist Fandey Mashimba works with a tiny population of a towering species called Karomia gigas. These trees, with their large oval leaves and distinctive, papery fruit, were thought to have gone extinct in the 1980s, but around six of them were discovered in 2011 by botanists from the University of Dar es Salaam. Protecting the habitat isn’t enough, because a fungus destroys their immature fruit. Mashimba, who oversees seed production for Tanzania’s Forest Service Agency, tries to whisk the fruit away before the fungus infects them, to sterilize and multiply the seeds for planting.Mashimba and his colleagues tried germinating hundreds of K. gigas seeds. The result: just three treasured plants, which Mashimba monitors through his office window as their giant leaves wave in the breeze. In 2018, the forestry service also dispatched 6,000 fruit to the Missouri Botanical Garden in St Louis. There, botanist Roy Gereau oversaw the extraction and cultivation of 24,000 seeds. The seeds produced only 30 plants. Last year, one sapling unfurled a small, pale purple flower, which perished within a day. When two trees flower simultaneously, botanists will attempt cross-pollination.

One of the 30 K. gigas plants at Missouri Botanical Garden flowered for a single day last year.Credit: Cassidy Moody/Missouri Botanical Garden

Mashimba is lucky in one respect: at least K. gigas produces seeds. Some trees produce none because their pollinators are gone; sometimes only one sex of a tree remains. For instance, most of the surviving specimens of the catkin yew (Amentotaxus argotaenia) in southern China are male. After a global search, a single female was discovered in the Royal Botanic Garden Edinburgh, UK; scientists there dispatched cuttings for planting near the surviving males. When they flower, reproduction can begin, says Gunter Fischer, a restoration ecologist at Missouri Botanical Garden. But this could take 30 years.Even if scientists do manage to acquire seeds from trees that are near extinction, germinating them can be tricky. Some go into dormancy, a protective state that, depending on the species, might be broken only through heating, cooling or scarring. Natural dormancy can last for years. Scientists try to circumvent it by culturing the embryo — the small section of a plant seed that will become the roots and stems — in a process known as embryo rescue.Every trick in the book The lonesome palm in the Curepipe Botanic Gardens — elderly, damaged and spindly — has seed problems, germination problems and more. It has resisted multiple rescuers since the 1980s. One obstacle is that the palm produces male and female flowers at different times, to avoid self-fertilization. Using a ladder and a brush, scientists override this process to collect, store and transfer pollen.It was the fruit of one such assisted-pollination project, each containing a single seed, that Sarasan carried back to Kew in 2006. He knew that lonesome palm seeds don’t grow if they are planted, so he used embryo rescue. With so few seeds, he felt there was no scope for experimenting with different culture media, so he made his best guess as to which blend to use.“I was so protective,” he says. “It was the responsibility, the excitement and also the fear of losing it.”The plantlets grew to 25 centimetres long. Then, one day, their fine white roots turned brown and they died, doubtless because of some nuance of the culture medium, he says.Other efforts have been derailed by mishap. In 2010, Kew horticultural scientist Carlos Magdalena negotiated to collect some freshly picked palm fruit while he was visiting Mauritius. Owing to a misunderstanding, two of the five fruit stored in a nearby fridge were eaten by a garden labourer who did not know their significance. Back at Kew, the seeds from the others failed to germinate.The failure rankles with Magdalena, who has a string of plant rescues to his name. As he roves the Kew greenhouses, steamy sanctuaries for plants that are bereft of a place in the wild, he sometimes feels he is all that stands between a species and its permanent loss.José Luis Marcelo Peña knows how he feels. In 2018, Marcelo Peña, a taxonomist at the National University of Jaén in Peru, was trekking through a steep, parched forest in Peru’s Marañón valley when he discovered a tree with light green flowers: Pradosia argentea, thought to be extinct.“It was a unique happiness that cannot be described,” says Marcelo Peña. Surveys yielded 200 more trees in the area, all of which were imminently threatened by agriculture.COVID-19 lockdowns began just as he attempted to save them. Without university facilities, but with remote help from the BGCI, he extracted 400 seeds from the purple fruit at home. More than 60 germinated: 20 survived. The following year, he tried again using fresh seeds, but a fungus got them all.As he finishes his story, he removes his glasses to wipe tears away. “It’s a big responsibility,” he says. And even with 20 little successes in the nursery, Marcelo Peña is concerned about the next step — reintroduction to the wild. Local people were unaware of P. argentea until recently, he says. They now support protecting the remaining trees — but they also need space to farm, which could put those survivors at risk.Back to the wildThriving in the wild is a distant dream for K. gigas, too. Tanzania’s forest agency and its partners are developing seed-propagation sites and nurseries for the species. But its future is uncertain, mostly because new trees could succumb to the same mysterious fungus.“We might have to content ourselves with saying, well, we have these lovely creatures in the zoo,” says Gereau.

A project at Missouri Botanical Garden produced 30 K. gigas plants.Credit: Cassidy Moody/Missouri Botanical Garden

Reintroductions can be spectacularly successful, however. The BGCI highlights a project on Malawi’s Mount Mulanje, the only natural home of the cypress Widdringtonia whytei. In 2019, just seven mature trees remained, the others victims of illegal felling. By 2022, thanks to a collaboration with Malawi’s Forestry Research Institute and local people, the slopes are alive again with 500,000 seedlings, and many locals now make a living through this endeavour.Propagation itself turned out to be fairly simple, says Smith. In Mauritius, by contrast, ecologists have a tougher task. The Mauritian Wildlife Foundation, with help from botanists elsewhere, is attempting to save multiple critically endangered species at once, but success at propagation varies widely. There have been some dramatic restorations, including of some species from which only a single tree remained. But the lonesome palm, now part of this project, continues to resist.

Scientists affix protective netting around hand-pollinated flowers on H. amaricaulis in Mauritius.Credit: Atmah Toocaram

A fourth attempt has begun. Nets hang around the tree to catch the male flowers and store their pollen for hand fertilization when the female flowers appear. In France, botanist Stéphane Buord at the National Botanical Conservatory of Brest hopes to overcome the problem that faced Sarasan — too few seeds — by tapping into the large quantities of seeds produced by Hyophorbe vaughanii, a close Mauritian relative of the lonesome palm. He and his team have spent years working out a complex technical protocol that coaxes its embryos into rooted seedlings that survive outside a test tube. Now he is waiting to try this approach on the seeds of the lonesome palm.If he succeeds, the palm might eventually be reintroduced into a national park or into the wild. Kersley Pynee, a conservation scientist at the Mauritius National Parks and Conservation Service, has reintroduced other trees and shrubs and says it is an uphill struggle. Plants can fall victim to fungi, pests and other assailants. After one recent planting of 1,000 seedlings of the flowering shrub Nesocodon mauritianus, just 5 now remain, he says.This is to be expected, says Smith. In nature, trees produce vast quantities of seeds, of which only a fraction germinate and survive because of natural dangers such as infestations, fire or competition for light or nutrients.Tree museumThe Global Trees Campaign has so far planted out hundreds of thousands of seedlings from 300 threatened tree species. But for trees that can no longer survive in the wild, the only other options are to keep a specimen in a living collection, or to store its seeds in a bank.One target of the 2011 Global Strategy for Plant Conservation, part of the Convention on Biological Diversity, was to conserve at least 75% of threatened plants in living collections or seed banks by 2020 — a goal that has not been met. What’s more, simply drying and freezing seeds doesn’t always work. Technologies such as cryopreservation — fast freezing at ultra-low temperatures — could offer an alternative, although it is expensive and impractical for many countries. And in 2018, conservationists warned4 that the seeds of one-third of tree species cannot be banked, largely because they don’t survive drying.Smith rejects this bleak diagnosis. Between seed banks, cryopreservation, nurseries, botanic gardens and arboreta, there are plenty of options to “buy us time”, he says.One trend that could help is mass tree-planting, in which governments and corporations plant trees to sequester carbon to meet emissions targets. Done badly, as many of these projects are, mass plantings can destroy biodiversity. Done well, they could rescue many species, says Smith. “This is a bandwagon we really need to jump on.”

This specimen of Encephalartos woodii, found in South Africa, was relocated in the late 1800s to the Royal Botanic Gardens at Kew, near London. It is the only one of this species to ever have been found in the wild.Credit: Andrew McRobb/RBG Kew

To help boost the usefulness of such projects to biodiversity, the BGCI and its partners have drawn up a certification programme for tree-planting projects called the Global Biodiversity Standard.Species conservation could also piggyback on the growing ecosystem-restoration movement. There are now more than 100,000 of these projects globally, helping ecosystems to capture carbon and provide essential services.Smith argues that including native species strengthens such projects. But restoration ecologists are often more concerned with overall function than with individual species, says Curt Meine, a historian of ecology at the Aldo Leopold Foundation in Baraboo, Wisconsin. And they want ecosystems to provide multiple services to humans, including sustainable livelihoods. Some acknowledge that tree conservation should have a place. “I do think it’s important work and we could learn a lot,” says Robin Chazdon, a restoration ecologist at the University of Connecticut in Storrs.But there are more threatened tree species than there are restoration projects to absorb them. “It’s not going to be the way of protecting all of those tree species,” she says.Some ecologists have deeper concerns. Bridgewater says that the efforts of conservationists and of restoration ecologists don’t factor in climate change.“They all in the end assume that nothing is going to be changing,” he says. But many trees, and whole ecosystems, just won’t survive in their current ranges, he says.“You could save every tree species but it will not be what people think — it will be in botanical gardens and larger managed conservation areas, and planting where it’s suitable for survival, not where it’s currently growing.”But the tree saviours are driven by something visceral: panic at the permanent loss of the rich, unique, irreplaceable and often-undeciphered identity of each species.“I don’t feel I am, as a humble human, here for a few decades on this planet, authorized to just cut off millions of years of evolutionary history,” says Vié. “Every species has a value.” More

Our study method includes the following stages: (1) framing the investigation problem, (2) examining the literature, (3) developing and verifying two hypotheses, (4) collecting data, (5) the multiple criteria examination of 173 countries by means of the Degree of Project Utility and Investment Value Assessments (INVAR) method, (6) calculating correlations between 33 indicators and the success of 173 countries, (7) building 12 regression models, (8) compiling eight Maps (of which seven are CSS Maps) visualizing national success and sustainability, (9) spatial perspective analysis, and (10) performing integrated linear regression, multi-variant design and multiple criteria analysis of national policy alternatives, in order to identify rational decisions.This research is a quantitative study to examine the way national success affects 12 indicators of the three dimensions of sustainability in 173 countries, and uses the data from 2020, or the latest available.As investigation methods, our CSS Maps and Models can make it easier to study interdependencies between country success and sustainability. Supplementary Section 1, 4, and 5 presents our literature analysis which is carried out to gain deeper insights into our CSS Maps and Models, and to better understand their components in the worldwide research context.The following two core hypotheses have been proposed and verified for this research:

Hypothesis 1—The increasing success of a country is generally accompanied by increasing values for the three dimensions of sustainability indicators, and declines in these indicators lead to decreases in the country’s success. Improving some sustainability indicators tends to improve other sustainability indicators.

Hypothesis 2—Changes in the number of countries and their traditional key indicators system do not make a very significant difference to the relative national sustainability and success values. Likewise, the boundaries of the seven country clusters discussed in this research do not excessively depend on specific traditional key systems of indicators used in their analysis.

Along with different sets of national 17 success (Supplementary Table S1) and 12 sustainability (Supplementary Table S2) indicators, the INVAR method46 (Supplementary Section 2 and Fig. S1) was used to measure and map the success of the 173 countries selected as the focus for this research. The traditional statistical indicator systems defining country success and the three dimensions of sustainability are based on studies from various countries analyzed and combined. The INVAR method calculates an integrated criterion characterizing the overall success of the countries. This integrated criterion is directly proportional to the relative effect the values and weights of the given criteria make on the country’s success. The multiple-criteria INVAR analysis method has been applied to various countries, including Asian nations47, ex-Soviet states48, and a group of 169 countries49.This research used data from the framework of variables taken from various databases and websites, including Transparency International, Global Data, Eurostat-OECD, the World Bank, Knoema, the World Health Organization, Global Finance, Freedom House, Heritage, the Global Footprint Network, Socioeconomic Data and Applications Center, Our World in Data, Climate Change Knowledge Portal (World Bank Group), and The Institute for Economics and Peace, as well as global and national statistics and publications. All 173 countries analyzed in this article are listed in matrices, along with their 17 detailed success (Supplementary Table S1) and 12 sustainability (Supplementary Table S2) indicators (systems of indicators, their numbering, values, and weights). The INVAR method46 was applied to perform multiple criteria analysis of the 173 countries, and the results are presented in Supplementary Table S1 and Figs. 2, 3, 4 and 5. We use equal and different weights of 17 indicators to calculate the deviation of priorities for the 173 countries, which stands at 5.34% (Supplementary Section 2 and Fig. S2).Along with different sets of 12 national sustainability and 17 success indicators, the INVAR method46 was used to measure and map the success of the 173 countries selected as the focus of this research. The traditional statistical indicator systems defining country success and the three dimensions of sustainability are based on studies from various countries analyzed and combined. The INVAR method calculates an integrated criterion characterizing the overall success of the countries. This integrated criterion is directly proportional to the relative effect the values and weights of the given criteria make on the country’s success.Supplementary Table S3 shows the correlations between all measures determined by analyzing 173 countries. Supplementary Table S4 reveals the correlation coefficient matrix of the 17 success criteria for each of the 173 countries analyzed in this survey.Along the vertical axis y we analyze seven sustainability indicators, and along the horizontal axis x we analyze the success and priority indicators (9 CSS Map dimensions). The median correlation between the survival versus self-expression values and the nine CSS Map dimensions (the x-axis and y-axis) is moderate, whereas the median correlation between the traditional versus secular–rational values and the nine CSS Map dimensions is strong (Fig. 1).Tables S5-S8 show the descriptive statistics of 12 CSS Models (Supplementary Section 3). Supplementary Table S8 shows the extent to which a 1% increase or decrease in success of country’s features can push sustainability indicators up or down, expressed as a percentage. Supplementary Table S8 also shows the degree to which the percentage changes of success or the values of country’s features explain or fail to explain the dispersion of sustainability indicators. These CSS Models (Supplementary Section 3) show that when a country’s success increases by 1%, its 12 indicators related to the three dimensions of sustainability improve by on average 0.85% (Supplementary Table S8). Furthermore, the 17 variables of country success used in the CSS Models explain 80.8% on average of the dispersion of the three dimensions of sustainability and 98.2% of the dispersion of the country success variable (Supplementary Table S8).An increase of 1% in a country’s success is accompanied by a 0.39% average increase in its social and environmental (0.84% on average) sustainability indicators (Supplementary Table S8). On average, the CSS Sustainability Models explain 76.3% of the dispersions among the environmental sustainability indicators, 83.4% of the dispersions among the social sustainability indicators, and 94.5% of the dispersion among economic (i.e. the gross national income per capita) sustainability indicators (Supplementary Table S8).The study produced the eight Maps (of which seven are CSS Maps) of the World based on an analysis of 99–150 countries (the 2020 Inglehart–Welzel Cultural Map of the World focused on 103 analogical CSS Maps countries). The two dimensions of country success on the CSS Maps are represented in a system of 17 variables (Supplementary Table S1). When a country’s success grows, its performance related to the three dimensions of sustainable development increases as well, and the eight Maps (of which seven are CSS Maps) clearly illustrate this relationship (Figs. 2, 3, 4 and 5). The CSS Maps of the World developed as part of this study are described in Supplementary Section 5.Studies from various countries and our research suggest that country success and their features (x-axis) and sustainability indicators (y-axis) are generally strongly interrelated, and move in the same direction over time. This means that successful countries also perform better on sustainability dimensions.Stage 9 involved analysis of the spatial perspective research in place for explaining and predicting globally recognised physical, spatial, and human patterns in multiple ways. We apply 12 CSS Models, alternative design and multi-criteria analysis methods for spatial perspective analysis (Supplementary Section 4).The following additional two research objectives were set: (1) to determine the impact of a country’s success factors on sustainability metrics, and (2) to offer stakeholders recommendations regarding the strategies for improving sustainability indicators. The ways to improve sustainability indicators are determined by analysing 17 dependent variables (the main paper section “Practical applications and implications”, Table S9). As previously mentioned, in stage 10, national policy options have been examined by means of integrated linear regression, multi-variant design and multiple criteria analysis to identify rational decisions. Analysis of multiple alternative options and their detailed indicators, with a consideration of the existing state of the micro, meso, and macro environment, can ensure rational country success and sustainability. Below, a brief analysis of several best global practice examples of ways to identify rational policy, activities, and strategy follows. The examples presented below suggest that multiple possible alternatives must be designed, assessed against a system of micro, meso and macro indicators, and the most effective options selected to make countries more sustainable. In Isham and Jackson’s14 opinion, materialistic lifestyles and values have been associated with adverse effects on human health as well as having detrimental effects on our planet. Therefore, activities and lifestyles should be identified that promote human well-being, yet which at the same time protect ecological security. Isham and Jackson14 identify optimal activities (arts and crafts, reading, sports, meditating) with high levels of human well-being and low environmental costs. It is important to estimate pollution impacts on health in order to come up with the right policies for better health outcomes. Yet, the task is challenging because economic activity can lead to worse pollution, but can also improve health outcomes in its own right37. Humidity, temperature, dispersal by the wind, and other environmental factors contribute to pollution levels. Certain fine particulates can stay in the atmosphere for days, and travel long distances to be inhaled in places far away from the source, even in other continents. Local conditions must be reflected in emissions-control policies, and the global flows of air pollutants must be taken into account6. The explanation for the phenomenon of demographic transition could be improved public health in developed countries which results in a move toward a slower life strategy38. Studies show that children from wealthier backgrounds undergo puberty later than those from poor socio-economic backgrounds. Early puberty can lead to a variety of health problems and a shorter life. By the early adult years, the effects of exposure to trauma, post-traumatic stress disorder, and other conditions can become apparent in the form of diseases related to aging9. Education is a very important factor in economic growth, and is also strongly related to health. In addition to health benefits, substantial increases in education, especially of women, and shrinking gender gaps have an important effect on the roles and status of women in society36.The INVAR method, statistical analysis, and the CSS Maps and Models can help generate multiple policy recommendations for various stakeholders. The possibilities are as follows:

To create alternatives for ways to develop country success and sustainability, by performing countries’ multiple criteria and statistical analysis and identifying decisions that would be rational;

to perform quantitative and qualitative analysis of the existing data and to interpret it. The results obtained this way would prompt automatic recommendations designed for different stakeholders on ways to improve country sustainability. More

Smouse, P. E. & Peakall, R. O. D. Spatial autocorrelation analysis of individual multiallele and multilocus genetic structure. Heredity 82(5), 561–573 (1999).PubMed 
Article 

Google Scholar 
Liu, J. X., Gao, T. X., Wu, S. F. & Zhang, Y. P. Pleistocene isolation in the Northwestern Pacific marginal seas and limited dispersal in a marine fish, Chelon haematocheilus (Temminck & Schlegel, 1845). Mol. Ecol. 16(2), 275–288 (2007).CAS 
PubMed 
Article 

Google Scholar 
Ding, S., Mishra, M., Wu, H., Liang, S. & Miyamoto, M. M. Characterization of hybridization within a secondary contact region of the inshore fish, Bostrychus sinensis, in the East China Sea. Heredity 120(1), 51–62 (2018).CAS 
PubMed 
Article 

Google Scholar 
Ashrafzadeh, M. R. et al. Assessing the origin, genetic structure and demographic history of the common pheasant (Phasianus colchicus) in the introduced European range. Sci. Rep. 11(1), 1–14 (2021).Article 
CAS 

Google Scholar 
Caccavo, J. A. et al. Along-shelf connectivity and circumpolar gene flow in Antarctic silverfish (Pleuragramma antarctica). Sci. Rep. 8(1), 1–16 (2018).Article 
CAS 

Google Scholar 
Otwoma, L. M., Reuter, H., Timm, J. & Meyer, A. Genetic connectivity in a herbivorous coral reef fish (Acanthurus leucosternon Bennet, 1833) in the Eastern African region. Hydrobiologia 806(1), 237–250 (2018).Article 

Google Scholar 
Li, H., Lin, H., Li, J. & Ding, S. Phylogeography of the Chinese beard eel, Cirrhimuraena chinensis Kaup, inferred from mitochondrial DNA: A range expansion after the last glacial maximum. Int. J. Mol. Sci. 15(8), 13564–13577 (2014).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Gao, B., Song, N., Li, Z., Gao, T. & Liu, L. Population genetic structure of Nuchequula mannusella (Perciformes: Leiognathidae) population in the Southern Coast of China inferred from complete sequence of mtDNA Cyt b gene. Pak. J. Zool. 51(4), 1527–1535 (2019).Article 

Google Scholar 
Qiu, F., Li, H., Lin, H., Ding, S. & Miyamoto, M. M. Phylogeography of the inshore fish, Bostrychus sinensis, along the Pacific coastline of China. Mol. Phylogenet. Evol. 96, 112–117 (2016).PubMed 
Article 

Google Scholar 
Gu, S. et al. Genetic diversity and population structure of cutlassfish (Lepturacanthus savala) along the coast of mainland China, as inferred by mitochondrial and microsatellite DNA markers. Reg. Stud. Mar. Sci. 43, 101702 (2021).
Google Scholar 
Liu, Q. et al. Genetic variation and population genetic structure of the large yellow croaker (Larimichthys crocea) based on genome-wide single nucleotide polymorphisms in farmed and wild populations. Fish. Res. 232, 105718 (2020).Article 

Google Scholar 
Song, P. et al. Genetic characteristics of yellow seabream Acanthopagrus latus (Houttuyn, 1782) (Teleostei: Sparidae) after stock enhancement in southeastern China coastal waters. Reg. Stud. Mar. Sci. 48, 102065 (2021).
Google Scholar 
Ward, R. D. Genetics in fisheries management. Hydrobiologia 420, 191–201 (2000).CAS 
Article 

Google Scholar 
Liu, X., Guo, Y., Wang, Z. & Liu, C. The complete mitochondrial genome sequence of Trichiurus nanhaiensis (Perciformes: Trichiuridae). Mitochondrial DNA 24(5), 516–517 (2013).CAS 
PubMed 
Article 

Google Scholar 
Wang, H. Y., Dong, C. A. & Lin, H. C. DNA barcoding of fisheries catch to reveal composition and distribution of cutlassfishes along the Taiwan coast. Fish. Res. 187, 103–109 (2017).Article 

Google Scholar 
Guo, Y. S., Liu, X. M., Wang, Z. D., Lu, H. S. & Liu, C. W. Isolation and characterization of microsatellite DNA loci from Naihai cutlassfish (Trichiurus nanhaiensis). J. Genet. 93(1), 109–112 (2014).Article 

Google Scholar 
Kwok, K. Y. & Ni, I. H. Reproduction of cutlassfishes Trichiurus spp. from the South China Sea. Mar. Ecol. Prog. Ser. 176, 39–47 (1999).ADS 
Article 

Google Scholar 
He, L. et al. Demographic response of cutlassfish (Trichiurus japonicus and T. nanhaiensis) to fluctuating palaeo-climate and regional oceanographic conditions in the China seas. Sci. Rep. 4(1), 1–10 (2014).
Google Scholar 
Lin, H. C., Tsai, C. J. & Wang, H. Y. Variation in global distribution, population structures, and demographic history for four Trichiurus cutlassfishes. PeerJ 9, e12639 (2021).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Funk, W. C., McKay, J. K., Hohenlohe, P. A. & Allendorf, F. W. Harnessing genomics for delineating conservation units. Trends Ecol. Evol. 27(9), 489–496 (2012).PubMed 
PubMed Central 
Article 

Google Scholar 
Tautz, D. Hypervariability of simple sequences as a general source for polymorphic DNA markers. Nucleic Acids Res. 17, 6463–6471 (1989).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Templeton, A. R. The, “Eve” hypotheses: A genetic critique and reanalysis. Am. Anthropol. 95, 51–72 (1993).Article 

Google Scholar 
Van Oosterhout, C., Hutchinson, W. F., Wills, D. P. & Shipley, P. MICRO-CHECKER: Software for identifying and correcting genotyping errors in microsatellite data. Mol. Ecol. Notes 4(3), 535–538 (2004).Article 
CAS 

Google Scholar 
Frankham, R. Challenges and opportunities of genetic approaches to biological conservation. Biol. Conserv. 143(9), 1919–1927 (2010).Article 

Google Scholar 
Sun, P., Shi, Z., Yin, F. & Peng, S. Population genetic structure and demographic history of Pampus argenteus in the Indo-West Pacific inferred from mitochondrial cytochrome b sequences. Biochem. Syst. Ecol. 43, 54–63 (2012).CAS 
Article 

Google Scholar 
Liu, S. Y. V. et al. Genetic stock structure of Terapon jarbua in Taiwanese waters. Mar. Coast. Fish. 7(1), 464–473 (2015).Article 

Google Scholar 
Song, C. Y., Sun, Z. C., Gao, T. X. & Song, N. Structure analysis of mitochondrial DNA control region sequences and its applications for the study of population genetic diversity of Acanthogobius ommaturus. Russ. J. Mar. Biol. 46(4), 292–301 (2020).CAS 
Article 

Google Scholar 
Yan, Y. R. et al. Cryptic diversity of the spotted scat Scatophagus argus (Perciformes: Scatophagidae) in the South China Sea: Pre-or post-production isolation. Mar. Freshw. Res. 71(12), 1640–1650 (2020).Article 

Google Scholar 
Hartl, D. L., Clark, A. G., & Clark, A. G. Principles of Population Genetics, vol. 116 (Sinauer Associates, 1997).Wu, R. et al. Study on the nomenclature and taxonomie status of hairtail Trichiurus japonicus from the Chinese coastal waters. Genom. Appl. Biol. 37(9), 3782–3791 (2018).
Google Scholar 
Xu, D. et al. Genetic diversity and population differentiation in the yellow drum Nibea albiflora along the coast of the China Sea. Mar. Biol. Res. 13(4), 456–462 (2017).Article 

Google Scholar 
Wang, W. et al. Genetic diversity and population structure analysis of Lateolabrax maculatus from Chinese coastal waters using polymorphic microsatellite markers. Sci. Rep. 11(1), 1–11 (2021).Article 
CAS 

Google Scholar 
Cheng, Q., Chen, W. & Ma, L. Genetic diversity and population structure of small yellow croaker (Larimichthys polyactis) in the Yellow and East China seas based on microsatellites. Aquat. Living Resour. 32, 16 (2019).Article 

Google Scholar 
DeWoody, J. A. & Avise, J. C. Microsatellite variation in marine, freshwater and anadromous fishes compared with other animals. J. Fish Biol. 56(3), 461–473 (2000).CAS 
Article 

Google Scholar 
Song, N., Yin, L., Sun, D., Zhao, L. & Gao, T. Fine-scale population structure of Collichtys lucidus populations inferred from microsatellite markers. J. Appl. Ichthyol. 35(3), 709–718 (2019).Article 

Google Scholar 
Yin, W. et al. Species delimitation and historical biogeography in the genus Helice (Brachyura: Varunidae) in the Northwestern Pacific. Zool. Sci. 26(7), 467–475 (2009).CAS 
Article 

Google Scholar 
Hewitt, G. The genetic legacy of the Quaternary ice ages. Nature 405(6789), 907–913 (2000).ADS 
CAS 
PubMed 
Article 

Google Scholar 
Yi, M. R. et al. Genetic structure and diversity of the yellowbelly threadfin bream Nemipterus bathybius in the Northern South China Sea. Diversity 13(7), 324 (2021).CAS 
Article 

Google Scholar 
Chen, X., Wang, J. J., Ai, W. M., Chen, H. & Lin, H. D. Phylogeography and genetic population structure of the spadenose shark (Scoliodon macrorhynchos) from the Chinese coast. Mitochondrial DNA Part A 29(7), 1100–1107 (2018).Article 
CAS 

Google Scholar 
Huang, W. et al. Genetic diversity and large-scale connectivity of the scleractinian coral Porites lutea in the South China Sea. Coral Reefs 37(4), 1259–1271 (2018).ADS 
Article 

Google Scholar 
Hou, G. et al. Identification of eggs and spawning zones of hairtail fishes Trichiurus (Pisces: Trichiuridae) in Northern South China Sea, using DNA barcoding. Front. Environ. Sci. 9, 703029 (2021).Article 

Google Scholar 
Yamaguchi, K., Nakajima, M. & Taniguchi, N. Loss of genetic variation and increased population differentiation in geographically peripheral populations of Japanese char Salvelinus leucomaenis. Aquaculture 308, S20–S27 (2010).Article 

Google Scholar 
Neo, M. L., Liu, L. L., Huang, D. & Soong, K. Thriving populations with low genetic diversity in giant clam species, Tridacna maxima and Tridacna noae, at Dongsha Atoll, South China Sea. Reg. Stud. Mar. Sci. 24, 278–287 (2018).
Google Scholar 
Jeanmougin, F., Thompson, J., Gouy, M., Higgins, D. & Gibson, T. Multiple sequence alignment with Clustal X. Trends Biochem. Sci. 23, 403–405 (1998).CAS 
PubMed 
Article 

Google Scholar 
Librado, P. & Rozas, J. DnaSP v5: A software for comprehensive analysis of DNA polymorphism data. Bioinformatics 25(11), 1451–1452 (2009).CAS 
PubMed 
Article 

Google Scholar 
Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Mol. Biol. Evol. 35, 1547–1549 (2018).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Zhang, D. et al. PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Mol. Ecol. Resour. 20(1), 348–355 (2020).PubMed 
Article 

Google Scholar 
Excoffier, L. & Lischer, H. E. Arlequin suite ver 3.5: A new series of programs to perform population genetics analyses under Linux and Windows. Mol. Ecol. Resour. 10(3), 564–567 (2010).PubMed 
Article 

Google Scholar 
Tajima, F. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics 123, 585–595 (1989).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Fu, Y. X. Statistical tests of neutrality of mutations against population growth, hitchhiking and background selection. Genetics 147(2), 915–925 (1997).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Drummond, A. J., Suchard, M. A., Xie, D. & Rambaut, A. Bayesian phylogenetics with BEAUti and the BEAST 1.7. Mol. Biol. Evol. 29(8), 1969–1973 (2012).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Rambaut, A., Suchard, M. A., Xie, D., & Drummond, A. J. Tracer v1.6. (2014). http://tree.bio.ed.ac.uk/software/tracer/ (accessed 23 Jan 2018).Bohonak, A. J. IBD (isolation by distance): A program for analyses of isolation by distance. J. Hered. 93(2), 153–154 (2002).CAS 
PubMed 
Article 

Google Scholar 
Weir, B. S. & Cockerham, C. C. Estimating F-statistics for the analysis of population structure. Evolution 38, 1358–1370 (1984).CAS 
PubMed 

Google Scholar 
Goudet, J. FSTAT (version 2.9.3): A program to estimate and test gene diversities and fixation indices. www.unil.ch/izea/softwares/fstat.html (2001).Pritchard, J. K., Stephens, M. & Donnelly, P. Inference of population structure using multilocus genotype data. Genetics 155(2), 945–959 (2000).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Earl, D. A. & VonHoldt, B. M. STRUCTURE HARVESTER: A website and program for visualizing STRUCTURE output and implementing the Evanno method. Conserv. Genet. Resour. 4(2), 359–361 (2012).Article 

Google Scholar 
Peakall, R. O. D. & Smouse, P. E. GENALEX 6: Genetic analysis in Excel. Population genetic software for teaching and research. Mol. Ecol. Notes 6(1), 288–295 (2006).Article 

Google Scholar 
Jombart, T., Devillard, S. & Balloux, F. Discriminant analysis of principal components: A new method for the analysis of genetically structured populations. BMC Genet. 11(1), 1–15 (2010).Article 

Google Scholar 
R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2015).Cornuet, J. M. et al. DIYABC v2.0: A software to make approximate Bayesian computation inferences about population history using single nucleotide polymorphism, DNA sequence and microsatellite data. Bioinformatics 30(8), 1187–1189 (2014).CAS 
PubMed 
Article 

Google Scholar 
Cornuet, J. M., Ravigne, V. & Estoup, A. Inference on population history and model checking using DNA sequence and microsatellite data with the software DIYABC (v1.0). BMC Bioinform. 11, 401 (2010).Article 
CAS 

Google Scholar  More

Highland lakes in southwestern China supply water to more than 1.4 billion people. Increasingly subject to eutrophication, biodiversity loss, drought and pollution, the lakes urgently need integrated management by government, community stakeholders and scientists to guide development of watershed policy and address these challenges.
Competing Interests
The authors declare no competing interests. More

To date, many kinds of potential predator–prey interactions have been demonstrated between ticks and a wide range of animals, including birds, mammals, and arthropods such as beetles, spiders, and ants1,2,3. Of those, the ants were indicated to be one of the most effective tick predators4,5. More than 27 ant species of 17 genera (Anoplolepis, Aphaenogaster, Camponotus, Crematogaster, Ectatomma, Formica, Iridomyrmex, Meranoplus, Monomorium, Myrmica, Notoncus, Pheidole, Pogonomyrmex, Polyrhachis, Rhytidoponera, Solenopsis, and Tapinoma) have been reported to be effective on different tick species (Amblyomma spp., Boophilus spp., Ornithodoros spp., Ixodes spp., Argas spp., Aponomma hydrosauri, Rhipicephalus appendiculatus, Otobius megnini, and Dermacentor variabilis)2,6,7,8. It is known that ants and other predators can affect ticks in a consumptive or nonconsumptive / behavioral manner and, as a result, may reduce the abundance of ticks in the overlap ranges8,9,10. However, the effects of ants on ticks are closely related to the ant species and the species, developmental stages, and physiological status of the ticks, and as a consequence, the impact of ants on ticks can exhibit fairly high variability2,8. Furthermore, there is no sufficient data on the factors determining the tick-ant relationship11,12.Ant predation has been examined in all developmental stages of ticks, but the proportion of the studies based on the eggs is relatively low compared to the other stages2. In an egg-based study, the eggs of O. megnini, the spinose ear tick, were supplied to five different ant species, and of those, Tapinoma melanocephalum was the only species that fed on the eggs7. Conflicting results have been reported from the studies13,14 carried out to determine the predatory effects of ant species Pheidole megacephala on the eggs of Boophilus (Rhipicephalus) microplus14. Rhipicephalus sanguineus was demonstrated to secrete an acarine allomone when attacked by fire ants, Solenopsis invicta15. This allomone-based ant deterrence is known to protect ticks from being eliminated within the sympatric range. The eggs, intact and cracked, of tick species Amblyomma americanum were not attacked by S. invicta, and it was interpreted that this deterrence might be related to the possible presence of the allomone within the eggs12.This study was carried out to determine whether the ant species Lasius alienus (Förster, 1850) (Hymenoptera: Formicidae) has any predatory effect on the eggs of tick species Hyalomma marginatum, H. excavatum, and Rhipicephalus bursa, and if the tick egg wax has any protective properties against possible predation. Ticks lay eggs (each 50–100 µg in weight and 0.5–1 mm in length) with a wax coat 0.5–2.0 µm thick, which is secreted by the female-tick-specific glands and organs such as the Gené’s16,17. Different molecules have been detected in the wax, such as alkanes, fatty acids, steroids, alcohols, and some specific proteins and lipoproteins18,19,20. However, detailed data on the wax content, especially its bioactive components, are not yet available20,21. As for the function of the wax, it has been reported that it reduces water loss, waterproofs the eggs, ensures the proper gas exchange between the eggs and air and holds the eggs together16,18. The wax also provides protection against chemical and physical factors such as cold, heat, proteinase K and pronase, or microbial agents including bacteria, fungi, viruses, and protozoa19,20,21,22,23,24,25.Lasius alienus is one of the most abundant ant species in the Western Palearctic26. Its distribution area can range from natural open habitats, light forests, and forest edges to urbanized areas such as wooded residential areas and gardens. The nests can be encountered mostly in the soil, under stones, or other substances, and the nest densities may reach up to 10–50 nests/100 m2 in some endemic territories. The number of workers (2­4 mm in length) in a colony can be more than 10,000. In the active periods in hot and warm months, workers establish foraging trails on the ground, in trees, and even in dwellings for food27,28. Lasius alienus was reported to gather plant nectar, honeydew secreted by aphids and to consume both dead and small living arthropods28,29,30. However, there is no data in the literature on whether this or any other species in the Lasius genera have a predatory interaction with ticks. Furthermore, the only definitive association of L. alienus with predation on Acarina has been established recently with Dermanyssus gallinae (poultry red mite)31.Considering the fact that the workers of L. alienus can forage effectively in many different places within the wide range distribution area27,28,31, it seems quite possible that several tick species encounter this ant species in their habitat32. This ant can feed by scavenging and predating small insects, and it meets their protein needs by hunting large numbers of small invertebrates, especially during larval feeding periods using the central foraging strategy29. Whichever invertebrates are abundant in their environment, the ants undoubtedly tend to consume more of them, especially if they are easy to hunt and transport27,28,29. Engorged large female ixodid ticks (around 1–1.5 cm depending on the species) lay a single batch of a large number of eggs (hundreds or thousands depending on the species and feeding levels) for several days or weeks at the hiding points such as cracks, crevices, and spaces under stones or various objects on the ground21,33 that the ant can easily reach due to its small size. In ixodid ticks, the female ticks lay eggs at once and die. The next generation continues entirely through the eggs21. Referring to these data, it seems hypothetically possible that any level of predation of L. alienus on the eggs, which are immobile and easy to reach and carry for the ants, can have a direct effect on the tick community in the overlap ranges. At this point, of course, whether the natural distribution areas of the ant and tick species overlap and, potentially, whether there is a possible evolutionary background between them are of the expected determining factors in a possible predation34.The nests of L. alienus can be seen almost everywhere in the soil in its ranging territories, however, the density increases from dry steppe, open pasture and bushlands to cultivated areas, woodlands, and gardens29. In this study, three ixodid tick species were selected that are more or less different from each other in terms of biology, ecology, and therefore the probability of encountering L. alienus in their natural habitat. Of these, H. marginatum is a two-host tick, the immature stages (larvae and nymphs) prefer rabbits, hedgehogs and birds to feed, and the adults (male and female) use particularly cattle as host. The immature stages of mostly three-host tick H. excavatum, wild rodents are the preferable host, and a wide range of large wild animals, cattle and some other domestic animals can be the host for the adults. Both species are known as arid environment ticks, however, in accordance with their different host preferences as well, H. excavatum is more prevalent in the arid open fields, steppe, and bushlands32,35. Both immature and adult stages of two-host tick R. bursa use primarily domestic ruminants to feed. Although there is no detailed data on the natural dynamics of R. bursa, this species is suspected of having a kind of peri-farm natural dynamics32. More