Ecology

Soil physical–chemical properties and HMs concentrationsThe soil physical–chemical properties and HMs concentrations are summarized in Table 1. The soil mean pH value was 6.17 and ranged from 4.16 to 9.04 in different sites. The samples sites of level for pH ≤ 6.0(acidic soil), 6.0  7.5(alkaline soil) were 51.5%, 36.4% and 12.1%, respectively. The average content of TN, TP and TK were 1.33 g kg−1, 1.16 g kg−1 and 23.6 g kg−1, and ranged from 0.7 g kg−1 to 2.4 g kg−1, 0.22 g kg−1 to 20.8 g kg−1 and 10.3 g k g−1 to 28 g kg−1, respectively.The mean concentration of Cd, Hg, As, Pb, Cr, Cu, Ni and Zn were 0.221, 0.155, 9.76, 32.2, 91.9, 35.2, 37.1 and 108.8 mg kg−1. Except Cd, the average concentration of Hg, As, Pb, Cr, Cu, Ni and Zn exceeded 93.8%, 7.1%, 6.3%, 17.8%, 25.3%, 10.7%, and 32.4% the soil background values for Chengdu, respectively, which indicates that HMs are enriched to a certain extent in soil. The CV of the HMs in the agricultural soils increased in the order Ni(10.1%), Cr(10.9%), Pb(15.4%), As(21.4%), Cd(31.1%), Zn(57.8%), Hg(58.1%) and Cu(59.9%). The exceptionally high variability of Cu, Hg and Zn indicates that these metals differed greatly with respect to different sites, and the existence of abnormally high values is the main reason that the CV was high. It further indicates that Cu, Hg and Zn may be affected by external interference factors. The mean concentration of all HMs in soil were below the risk screening values for soil contamination (GB 15618-2018) (MEEC, 2018), however, the results showed that in 76, 1, 1, 6 and 2 of sample sites the level of Cd, Pb, Cr, Cu and Zn exceeded the risk screening values.Assessment of heavy metal(loid)s pollutionIndex of geo-accumulationThe index of geo-accumulation of HMs in the soil in the study area are shown in Fig. 2. In a descending order of magnitude of Igeo mean value, the eight elements were as follows: Hg(0.18)  > Zn(-0.22)  > Cu(-0.30)  > Cr(-0.36)  > Ni(-0.45)  > Pb(-0.51)  > As(-0.52)  > Cd(-0.82), indicating that the soil HMs Zn, Cu, Cr, Ni, Pb, As and Cd in the study area were generally in a no contamination according to the defined classes, while Hg was in uncontaminated to moderately contaminated.Figure 2Indexes of geo-accumulation, pollution indexes and potential ecological risk indexes of HMs in study aera. Circles at the top and bottom of box plots correspond to the maximum and minimum values, respectively. The square in the box plot is the average value. Horizontal lines at the top, middle, and bottom of the box plot correspond to 75% percentile, median, and 25% percentile, respectively.Full size imageThe Igeo values for Zn, Cu, Cr, Ni, Pb, As and Cd in more than 90% of samples were less than zero, only a few outliers in the soil were classified as moderately contaminated or worse. Which meaning point pollution at those sample sites. However, the Igeo values for Hg in 47.34%, 40.61%,10.66% and 1.40% of samples were belonged to  As  > Pb  > Ni  > Cu  > Hg  > Zn  > Cd. Cr, As and Pb were the largest contributors for both adults and children, accounting for 47.33% and 42.37%, 32.68% and 39.64%, 15.95% and 13.26%, respectively. Indicating that attention should be pain to the Cr, As and Pb elements due to their noncarcinogenic risk. Which was basically consistent with the research results of Bo et al.1 and Bao et al.32. Overall, The HMs of Cr, As and Pb were the main non-carcinogenic factors in soil in the study area, and the risk control of these elements should be strengthened.Carcinogenic risk assessmentThe CR of the HMs are shown in Table 5. Three exposure pathways were considered for Cd and Hg in our study. But As and Pb were considered carcinogenic by ingestion and inhalation, and Cr was considered carcinogenic through inhalation. The TCR mean values were 3.68 × 10−5 for adults and 6.27 × 10−5 for children, obviously were in the range 1 × 10−4 from 1 × 10−6, suggesting the TCR caused by HMs in the study area was acceptable on the whole, but it still exceeds the soil treatment threshold value 10−6. The CR average values through the three exposure pathways were CRing 3.65 × 10−5, CRinh 2.53 × 10−7 and CRderm 1.07 × 10−7 for adults, CRing 6.25 × 10−5, CRinh 1.12 × 10−7 and CRderm 4.50 × 10−8 for adults. Clearly the CRing was much larger than CRinh and CRderm for both adults and children, which indicates that oral ingestion is the major exposure pathway for CR. Which was consistent with the research results of Song et al.31 and Bo et al.1. For single HM, the CR value of Pb and Hg for adults were 2.72 × 10−5 and 8.6 × 10−6, and Pb, Hg and Cd for children were 4.63 × 10−5, 1.48 × 10−5 and 1.36 × 10−6, respectively, within acceptable criterion. Overall, the longterm health effects for adults and children are not serious at current single HM level.In this study, the total contents of HMs in the soils were used to assess health risk, the bioavailability of HMs were not considered, which may have caused the assessment results to be higher than the actual local situation1,31,42. In addition, because the parameters of health risk evaluation for children were set to be more sensitive than those for adults, the non-carcinogenic risk and carcinogenic risk for children were higher than those for adults2,43,44. However, those risks were at acceptable or negligible levels. Therefore, the study area is suitable for safe and clean production of hotbed chives. More

Fisher, M. C. et al. Threats posed by the fungal kingdom to humans, wildlife, and agriculture. MBio 11, e00449–20 (2020).PubMed 
PubMed Central 
Article 

Google Scholar 
Fisher, M. C., Hawkins, N. J., Sanglard, D. & Gurr, S. J. Worldwide emergence of resistance to antifungal drugs challenges human health and food security. Science 360, 739–742 (2018).CAS 
PubMed 
Article 

Google Scholar 
Nash, A. et al. MARDy: Mycology Antifungal Resistance Database. Bioinformatics 34, 3233–3234 (2018).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Ksiezopolska, E. et al. Narrow mutational signatures drive acquisition of multidrug resistance in the fungal pathogen Candida glabrata. Curr. Biol. 4, 5314–5326.e10 (2021).Article 
CAS 

Google Scholar 
Bryce Taylor, M. et al. yEvo: Experimental evolution in high school classrooms selects for novel mutations and epistatic interactions that impact clotrimazole resistance in S. cerevisiae. Preprint at bioRxiv https://doi.org/10.1101/2021.05.02.442375 (2021).Andersson, D. I. & Hughes, D. Antibiotic resistance and its cost: is it possible to reverse resistance? Nat. Rev. Microbiol. 8, 260–271 (2010).CAS 
PubMed 
Article 

Google Scholar 
Gerstein, A. C., Lo, D. S. & Otto, S. P. Parallel genetic changes and nonparallel gene-environment interactions characterize the evolution of drug resistance in yeast. Genetics 192, 241–252 (2012).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Yang, F. et al. The fitness costs and benefits of trisomy of each Candida albicans chromosome. Genetics 218, iyab056 (2021).PubMed 
PubMed Central 
Article 

Google Scholar 
Kanafani, Z. A. & Perfect, J. R. Antimicrobial resistance: resistance to antifungal agents: mechanisms and clinical impact. Clin. Infect. Dis. 46, 120–128 (2008).PubMed 
Article 

Google Scholar 
Iyer, K. R., Revie, N. M., Fu, C., Robbins, N. & Cowen, L. E. Treatment strategies for cryptococcal infection: challenges, advances and future outlook. Nat. Rev. Microbiol. 19, 454–466 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Longley, D. B., Harkin, D. P. & Johnston, P. G. 5-fluorouracil: mechanisms of action and clinical strategies. Nat. Rev. Cancer 3, 330–338 (2003).CAS 
PubMed 
Article 

Google Scholar 
Erbs, P., Exinger, F. & Jund, R. Characterization of the Saccharomyces cerevisiae FCY1 gene encoding cytosine deaminase and its homologue FCA1 of Candida albicans. Curr. Genet. 31, 1–6 (1997).CAS 
PubMed 
Article 

Google Scholar 
Wrenbeck, E. E., Azouz, L. R. & Whitehead, T. A. Single-mutation fitness landscapes for an enzyme on multiple substrates reveal specificity is globally encoded. Nat. Commun. 8, 15695 (2017).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Chen, J. Z., Fowler, D. M. & Tokuriki, N. Comprehensive exploration of the translocation, stability and substrate recognition requirements in VIM-2 lactamase. eLife 9, e56707 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Li, A., Acevedo-Rocha, C. G. & Reetz, M. T. Boosting the efficiency of site-saturation mutagenesis for a difficult-to-randomize gene by a two-step PCR strategy. Appl. Microbiol. Biotechnol. 102, 6095–6103 (2018).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Biot-Pelletier, D. & Martin, V. J. J. Seamless site-directed mutagenesis of the Saccharomyces cerevisiae genome using CRISPR-Cas9. J. Biol. Eng. 10, 6 (2016).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Dionne, U. et al. Protein context shapes the specificity of SH3 domain-mediated interactions in vivo. Nat. Commun. 12, 1597 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Eddy, A. A. Expulsion of uracil and thymine from the yeast Saccharomyces cerevisiae: contrasting responses to changes in the proton electrochemical gradient. Microbiology 143, 219–229 (1997).CAS 
PubMed 
Article 

Google Scholar 
Kurtz, J. E., Exinger, F., Erbs, P. & Jund, R. New insights into the pyrimidine salvage pathway of Saccharomyces cerevisiae: requirement of six genes for cytidine metabolism. Curr. Genet. 36, 130–136 (1999).CAS 
PubMed 
Article 

Google Scholar 
Fujimura, H. Growth inhibition of Saccharomyces cerevisiae by the immunosuppressant leflunomide is due to the inhibition of uracil uptake via Fur4p. Mol. Gen. Genet. 260, 102–107 (1998).CAS 
PubMed 
Article 

Google Scholar 
Després, P. C., Dubé, A. K., Nielly-Thibault, L., Yachie, N. & Landry, C. R. Double selection enhances the efficiency of Target-AID and Cas9-based genome editing in yeast. G3 8, 3163–3171 (2018).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Wang, J. et al. Role of glutamate 64 in the activation of the prodrug 5-fluorocytosine by yeast cytosine deaminase. Biochemistry 51, 475–486 (2012).CAS 
PubMed 
Article 

Google Scholar 
Ivankov, D. N., Finkelstein, A. V. & Kondrashov, F. A. A structural perspective of compensatory evolution. Curr. Opin. Struct. Biol. 26, 104–112 (2014).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Mayrose, I., Graur, D., Ben-Tal, N. & Pupko, T. Comparison of site-specific rate-inference methods for protein sequences: empirical Bayesian methods are superior. Mol. Biol. Evol. 21, 1781–1791 (2004).CAS 
PubMed 
Article 

Google Scholar 
Tarassov, K. An in vivo map of the yeast protein interactome. Science 320, 1465–1470 (2008).CAS 
PubMed 
Article 

Google Scholar 
Freschi, L., Torres-Quiroz, F., Dubé, A. K. & Landry, C. R. qPCA: a scalable assay to measure the perturbation of protein–protein interactions in living cells. Mol. Biosyst. 9, 36–43 (2013).CAS 
PubMed 
Article 

Google Scholar 
Chang, A. et al. BRENDA, the ELIXIR core data resource in 2021: new developments and updates. Nucleic Acids Res. 49, D498–D508 (2021).CAS 
PubMed 
Article 

Google Scholar 
Jumper, J. et al. Highly accurate protein structure prediction with AlphaFold. Nature 596, 583–589 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Mirdita, M. et al. ColabFold – Making protein folding accessible to all. Preprint at bioRxiv https://doi.org/10.1101/2021.08.15.456425 (2022).Pokusaeva, V. O. et al. An experimental assay of the interactions of amino acids from orthologous sequences shaping a complex fitness landscape. PLoS Genet. 15, e1008079 (2019).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Oliver, J. D. et al. F901318 represents a novel class of antifungal drug that inhibits dihydroorotate dehydrogenase. Proc. Natl Acad. Sci. USA 113, 12809–12814 (2016).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Hoenigl, M. et al. The antifungal pipeline: fosmanogepix, ibrexafungerp, olorofim, opelconazole, and rezafungin. Drugs https://doi.org/10.1007/s40265-021-01611-0 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Verweij, P. E., Te Dorsthorst, D. T. A., Janssen, W. H. P., Meis, J. F. G. M. & Mouton, J. W. In vitro activities at pH 5.0 and pH 7.0 and in vivo efficacy of flucytosine against Aspergillus fumigatus. Antimicrob. Agents Chemother. 52, 4483–4485 (2008).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Gsaller, F. et al. Mechanistic basis of pH-dependent 5-flucytosine resistance in Aspergillus fumigatus. Antimicrob. Agents Chemother. https://doi.org/10.1128/AAC.02593-17 (2018).Garland, T. Jr. Trade-offs. Curr. Biol. 24, R60–R61 (2014).CAS 
PubMed 
Article 

Google Scholar 
Chang, Y. C. et al. Moderate levels of 5-fluorocytosine cause the emergence of high frequency resistance in cryptococci. Nat. Commun. 12, 3418 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Billmyre, R. B., Applen Clancey, S., Li, L. X., Doering, T. L. & Heitman, J. 5-fluorocytosine resistance is associated with hypermutation and alterations in capsule biosynthesis in Cryptococcus. Nat. Commun. 11, 127 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Brachmann, C. B. et al. Designer deletion strains derived from Saccharomyces cerevisiae S288C: a useful set of strains and plasmids for PCR-mediated gene disruption and other applications. Yeast 14, 115–132 (1998).CAS 
PubMed 
Article 

Google Scholar 
Gietz, R. D. & Schiestl, R. H. High-efficiency yeast transformation using the LiAc/SS carrier DNA/PEG method. Nat. Protoc. 2, 31–34 (2007).CAS 
PubMed 
Article 

Google Scholar 
Janke, C. et al. A versatile toolbox for PCR-based tagging of yeast genes: new fluorescent proteins, more markers and promoter substitution cassettes. Yeast 21, 947–962 (2004).CAS 
PubMed 
Article 

Google Scholar 
Goldstein, A. L. & McCusker, J. H. Three new dominant drug resistance cassettes for gene disruption in Saccharomyces cerevisiae. Yeast 15, 1541–1553 (1999).CAS 
PubMed 
Article 

Google Scholar 
DeLuna, A., Springer, M., Kirschner, M. W. & Kishony, R. Need-based up-regulation of protein levels in response to deletion of their duplicate genes. PLoS Biol. 8, e1000347 (2010).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Casadaban, M. J. & Cohen, S. N. Analysis of gene control signals by DNA fusion and cloning in Escherichia coli. J. Mol. Biol. 138, 179–207 (1980).CAS 
PubMed 
Article 

Google Scholar 
Yachie, N. et al. Pooled-matrix protein interaction screens using barcode fusion genetics. Mol. Syst. Biol. 12, 863 (2016).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Andrews, S. FastQC: A quality control analysis tool for high throughput sequencing data (Babraham Bioinformatics, 2016); https://www.bioinformatics.babraham.ac.uk/projects/fastqc/Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Hunter, J. D. Matplotlib: a 2D graphics environment. Comput. Sci. Eng. 9, 90–95 (2007).Article 

Google Scholar 
Harris, C. R. et al. Array programming with NumPy. Nature 585, 357–362 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Reback, J. et al. pandas-dev/pandas: Pandas 1.3.4. Zenodo https://doi.org/10.5281/zenodo.5574486 (2021).Waskom, M. seaborn: statistical data visualization. J. Open Source Softw. 6, 3021 (2021).Article 

Google Scholar 
Virtanen, P. et al. SciPy 1.0: fundamental algorithms for scientific computing in Python. Nat. Methods 17, 261–272 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Masella, A. P., Bartram, A. K., Truszkowski, J. M., Brown, D. G. & Neufeld, J. D. PANDAseq: paired-end assembler for illumina sequences. BMC Bioinform. https://doi.org/10.1186/1471-2105-13-31 (2012).Rognes, T., Flouri, T., Nichols, B., Quince, C. & Mahé, F. VSEARCH: a versatile open source tool for metagenomics. PeerJ 4, e2584 (2016).PubMed 
PubMed Central 
Article 

Google Scholar 
Rice, P., Longden, L. & Bleasby, A. EMBOSS: the European Molecular Biology Open Software Suite. Trends Genet. https://doi.org/10.1016/S0168-9525(00)02024-2 (2000).Article 
PubMed 

Google Scholar 
Ryan, O. W., Poddar, S. & Cate, J. H. D. Crispr–cas9 genome engineering in Saccharomyces cerevisiae cells. Cold Spring Harb. Protoc. https://doi.org/10.1101/pdb.prot086827 (2016).Amberg, D. C., Burke, D. J. & Strathern, J. N. Methods in Yeast Genetics: A Cold Spring Harbor Laboratory Course Manual (CSHL Press, 2005).Ireton, G. C., Black, M. E. & Stoddard, B. L. The 1.14 A crystal structure of yeast cytosine deaminase: evolution of nucleotide salvage enzymes and implications for genetic chemotherapy. Structure 11, 961–972 (2003).CAS 
PubMed 
Article 

Google Scholar 
Schymkowitz, J. et al. The FoldX web server: an online force field. Nucleic Acids Res. 33, W382–W388 (2005).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Marchant, A. et al. The role of structural pleiotropy and regulatory evolution in the retention of heteromers of paralogs. eLife 8, e46754 (2019).PubMed 
PubMed Central 
Article 

Google Scholar 
Usmanova, D. R. et al. Self-consistency test reveals systematic bias in programs for prediction change of stability upon mutation. Bioinformatics 34, 3653–3658 (2018).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Howe, K. L. et al. Ensembl 2021. Nucleic Acids Res. 49, D884–D891 (2021).CAS 
PubMed 
Article 

Google Scholar 
Chorostecki, U., Molina, M., Pryszcz, L. P. & Gabaldón, T. MetaPhOrs 2.0: integrative, phylogeny-based inference of orthology and paralogy across the tree of life. Nucleic Acids Res. 48, W553–W557 (2020).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Byrne, K. P. & Wolfe, K. H. The Yeast Gene Order Browser: combining curated homology and syntenic context reveals gene fate in polyploid species. Genome Res. 15, 1456–1461 (2005).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Edgar, R. C. MUSCLE: a multiple sequence alignment method with reduced time and space complexity. BMC Bioinform. 5, 113 (2004).Article 
CAS 

Google Scholar 
Letunic, I. & Bork, P. Interactive Tree Of Life (iTOL) v5: an online tool for phylogenetic tree display and annotation. Nucleic Acids Res. 49, W293–W296 (2021).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Lõoke, M., Kristjuhan, K. & Kristjuhan, A. Extraction of genomic DNA from yeasts for PCR-based applications. Biotechniques 50, 325–328 (2011).PubMed 
PubMed Central 
Article 
CAS 

Google Scholar 
Schlecht, U., Miranda, M., Suresh, S., Davis, R. W. & St Onge, R. P. Multiplex assay for condition-dependent changes in protein-protein interactions. Proc. Natl Acad. Sci. USA 109, 9213–9218 (2012).CAS 
PubMed 
PubMed Central 
Article 

Google Scholar 
Diss, G. & Lehner, B. The genetic landscape of a physical interaction. eLife 7, e32472 (2018).PubMed 
PubMed Central 
Article 

Google Scholar 
Pettersen, E. F. et al. UCSF ChimeraX: structure visualization for researchers, educators, and developers. Protein Sci. 30, 70–82 (2021).CAS 
PubMed 
Article 

Google Scholar  More

Lucas, C. H. et al. Gelatinous zooplankton biomass in the global oceans: Geographic variation and environmental drivers. Glob. Ecol. Biogeogr. 23, 701–714. https://doi.org/10.1111/Geb.12169 (2014).Article 

Google Scholar 
Condon, R. H. et al. Recurrent jellyfish blooms are a consequence of global oscillations. Proc. Natl. Acad. Sci. USA 110, 1000–1005. https://doi.org/10.1073/pnas.1210920110 (2013).ADS 
Article 
PubMed 

Google Scholar 
Graham, W. M. et al. Linking human well-being and jellyfish: Ecosystem services, impacts, and societal responses. Front. Ecol. Environ. 12, 515–523. https://doi.org/10.1890/130298 (2014).Article 

Google Scholar 
Lucas, C. H., Gelcich, S. & Uye, S. I. Living with jellyfish: Management and adaptation strategies. In Jellyfish Blooms (eds Pitt, K. A. & Lucas, C. H.) 129–150 (Springer, 2014).Chapter 

Google Scholar 
De Donno, A. et al. Impact of stinging jellyfish proliferations along south Italian coasts: Human health hazards, treatment and social costs. Int. J. Environ. Res. Public Health 11, 2488–2503 (2014).PubMed 
PubMed Central 
Article 

Google Scholar 
Bosch-Belmar, M. et al. Consequences of stinging plankton blooms on finfish mariculture in the Mediterranean Sea. Front. Mar. Sci. 4, 240. https://doi.org/10.3389/fmars.2017.0024 (2017).Article 

Google Scholar 
Mayer, A. G. Medusae of the World: The Hydromedusae 132–498 (Carnegie institution of Washington, 1910).Book 

Google Scholar 
Kramp, P. L. Synopsis of the medusae of the world. J. Mar. Biol. Assoc. UK 40, 1–469 (1961).
Google Scholar 
Canepa, A. et al. Pelagia noctiluca in the Mediterranean Sea. In Jellyfish Blooms (eds Pitt, K. A. & Lucas, C. H.) 237–266 (Springer, 2014).Chapter 

Google Scholar 
Marambio, M. et al. Unfolding jellyfish bloom dynamics along the Mediterranean basin by transnational citizen science initiatives. Diversity 13, 274. https://doi.org/10.3390/d13060274 (2021).Article 

Google Scholar 
Mamish, S., Durgham, H. & Ikhtiyar, S. The first Pelagia noctiluca outbreak off the Syrian coast (the eastern Mediterranean Sea), five years after its first appearance. SSRG Int. J. Agric. Environ. Sci. 6, 72–75 (2019).
Google Scholar 
Daly Yahia, M. N. et al. Are outbreaks of Pelagia noctiluca (Forsskäl, 1775) more frequent in the Mediterranean Basin?. ICES Coop. Res. Rep. 300, 8–14 (2010).
Google Scholar 
Aissi, M., Touzri, C., Gueroun, S. K. M., Kefi-Daly Yahia, O. & Daly Yahia, M. N. Persistent occurrence and life cycle of Pelagia noctiluca in the channel of Bizerte (Northern Tunisia). Ecol. Environ. Conserv. 20, 1453–1460 (2014).
Google Scholar 
Kogovsĕk, T., Bogunović, B. & Malej, A. Recurrence of bloom forming scyphomedusae: Wavelet analysis of a 200-year time series. Hydrobiologia 645, 81–96 (2010).Article 
CAS 

Google Scholar 
Pestoric, B. et al. Scyphomedusae and ctenophora of the eastern adriatic: Historical overview and new data. Diversity 13, 186. https://doi.org/10.3390/d13050186 (2021).CAS 
Article 

Google Scholar 
UNEP (United Nations Environmental Programme). Workshop on Jellyfish Blooms in the Mediterranean, Athens (1984).UNEP (United Nations Environmental Programme). Jellyfish blooms in the Mediterranean Sea. Proceedings of II Workshop on Jellyfish in the Mediterranean Sea, Athens (1991).Goy, J., Morand, P. & Etienne, M. Long term fluctuations of Pelagia noctiluca (Cnidaria, Scyphomedusa) in the western Mediterranean. Sea Prediction by climatic variables. Deep-Sea Res. A 36, 269–279 (1989).ADS 
Article 

Google Scholar 
Bernard, P., Berline, L. & Gorsky, G. Long term (1981–2008) monitoring of the jellyfish Pelagia noctiluca (Cnidaria, Scyphozoa) on the French Mediterranean Coasts. J. Oceanogr. Res. Data 4, 1–10 (2011).
Google Scholar 
Brotz, L., Cheung, W. W. L., Kleisner, K., Pakhomov, E. & Pauly, D. Increasing jellyfish population: Trends in large marine ecosystems. Hydrobiologia 690, 3–20 (2012).Article 

Google Scholar 
Rosa, S., Pansera, M., Granata, A. & Guglielmo, L. Interannual variability, growth, reproduction and feeding of Pelagia noctiluca (Cnidaria: Scyphozoa) in the Straits of Messina (Central Mediterranean Sea): Linkages with temperature and diet. J. Mar. Syst. 111–112, 97–107 (2013).Article 

Google Scholar 
Aoutien, M., Bekkali, R., Nachit, D., Luan, K. & Mrhraoui, M. Predicting jellyfish strandings in the Moroccan North-West Mediterranean coastline. Eur. Sci. J. 15, 72–84. https://doi.org/10.19044/esj.2019.v15n2p72 (2019).Article 

Google Scholar 
Lynam, C. P., Hay, S. J. & Brierley, A. S. Interannual variability in abundance of North Sea jellyfish and links to the North Atlantic Oscillation. Limnol. Oceanogr. 49, 637–643 (2004).ADS 
Article 

Google Scholar 
Lynam, C. P. et al. Have jellyfish in the Irish Sea benefited from climate change and overfishing?. Glob. Change Biol. 17, 767–782 (2011).ADS 
Article 

Google Scholar 
Brodeur, R. D. et al. Rise and fall of jellyfish in the eastern Bering Sea in relation to climate regime shifts. Prog. Oceanogr. 77, 103–111 (2008).ADS 
Article 

Google Scholar 
Molinero, J. C. et al. Climate control on the longterm anomalous changes of zooplankton communities in the Northwestern Mediterranean. Glob. Change Biol. 14, 11–26 (2008).ADS 
Article 

Google Scholar 
Licandro, P. et al. A blooming jellyfish in the northeast Atlantic and Mediterranean. Biol. Let. 6, 688–691 (2010).CAS 
Article 

Google Scholar 
Ferraris, M. et al. Distribution of Pelagia noctiluca (Cnidaria, Scyphozoa) in the Ligurian Sea (NW Mediterranean Sea). J. Plankton Res. 34, 874–885 (2012).Article 

Google Scholar 
Malačič, V., Petelin, B. & Malej, A. Advection of the jellyfish Pelagia noctiluca (Scyphozoa) studied by the Lagrangian tracking of water mass in the climatic circulation of the Adriatic Sea. Geophys. Res. Abstr. 9, 02802 (2007).
Google Scholar 
Rubio, P. & Muñoz, J. M. Predicción estival del riesgo de blooms de Pelagia noctiluca (litoral central catalán). In Situaciones de riesgo climático en España (ed. Novau, J. C.) 281–287 (Instituto Pirenaico de Ecología, 1997).
Google Scholar 
Berline, L., Zakardjian, B., Molcard, A., Ourmieres, Y. & Guihou, K. Modeling jellyfish Pelagia noctiluca transport and stranding in the Ligurian Sea. Mar. Pollut. Bull. 70, 90–99 (2013).CAS 
PubMed 
Article 

Google Scholar 
Olds, A. D. et al. Quantifying the conservation value of seascape connectivity: A global synthesis. Glob. Ecol. Biogeogr. 25, 3–15 (2016).Article 

Google Scholar 
Vodopivec, M., Peliz, A. J. & Malej, A. Offshore marine constructions as propagators of moon jellyfish dispersal. Environ. Res. Lett. 12, 084003 (2017).ADS 
Article 

Google Scholar 
Chen, J. Z., Huang, S. L. & Han, Y. S. Impact of long-term habitat loss on the Japanese eel Anguilla japonica. Estuar. Coast. Shelf Sci. 151, 361–369 (2014).ADS 
CAS 
Article 

Google Scholar 
Fernandez-Arcaya, U. et al. Ecological role of submarine canyons and need for canyon conservation: A review. Front. Mar. Sci. 4, 5. https://doi.org/10.3389/fmars.2017.00005 (2017).Article 

Google Scholar 
Würtz, M. Towards a Mediterranean canyon inventory. Workshop (EBSAs), 7 to 11 April 2014, Málaga, Spain, 1–4 (2014).Sacchetti, F. Il ritorno di MeteoMedusa. Focus (Madison) 237, 92–94 (2012).
Google Scholar 
Benedetti-Cecchi, L. et al. Deterministic factors overwhelm stochastic environmental fluctuations as drivers of jellyfish outbreaks. PLoS ONE 10, e0141060. https://doi.org/10.1371/journal.pone.0141060 (2015).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Malej, A. & Malej, M. Population dynamics of the jellyfish Pelagia noctiluca (Forsskäl, 1775). In Proceedings of the 25th EMBS, Marine Eutrophication and Population Dynamics (ed. Colombo, G.A.) 215–219 (Olsen & Olsen, 1992).Rottini-Sandrini, L., Avian, M., Axiak, V. & Malej, A. The breeding period of Pelagia noctiluca (Scyphozoa, Semaeostomeae) in the Adriatic and central Mediterranean Sea. Nova Thalass. 6, 65–75 (1983).
Google Scholar 
Milisenda, G. et al. Reproductive and bloom patterns of Pelagia noctiluca in the Strait of Messina, Italy. Estuar. Coast. Shelf Sci. 201, 29–39. https://doi.org/10.1016/j.ecss.2016.01.002 (2018).ADS 
Article 

Google Scholar 
Magazzù, G. et al. Picoplankton: Contribution to phytoplankton production in the Strait of Messina. Mar. Ecol. 8, 21–31 (1987).ADS 
Article 

Google Scholar 
Guglielmo, L., Crescenti, N., Costanzo, G. & Zagami, G. Zooplankton and micronekton communities in the Straits of Messina. In The Straits of Messina ecosystem, present knowledge for an ecohydrodynamical approach. Proceedings of Symposium held in Messina, 4–6 April 1991, Messina (eds. Guglielmo, L., Manganaro, A. & De Domenico, E.) 247–270 (Dipartimento di Biologia Animale ed Ecologia, 1995).Guglielmo, L. et al. The Strait of Messina: A key area for Pelagia noctiluca (Cnidaria, Scyphozoa). In Jellyfish: Ecology, Distribution Patterns and Human Interactions (ed. Mariottini, G. L.) 71–90 (Nova Science Publishers Inc., 2017).
Google Scholar 
Astraldi, M. & Gasparini, G. P. The seasonal characteristics of the circulation in the Tyrrhenian Sea. In: Seasonal and Interannual Variability of the Western Mediterranean Sea, Coast. Estuar. Studies, Vol. 46, 115–134 (American Geophysical Union, 1994).Krivosheya, V. G. Water circulation and structure in the Tyrrhenian Sea. Oceanology 23, 166–171 (1983).
Google Scholar 
Millot, C. Circulation in the Western Mediterranean Sea. J. Mar. Syst. 20, 423–442. https://doi.org/10.1016/S0924-7963(98)00078-5 (1999).Article 

Google Scholar 
Vetrano, A., Napolitano, E., Iacono, R., Schroeder, K. & Gasparini, G. P. Tyrrhenian Sea circulation and water mass fluxes in spring 2004: Observations and model results. J. Geophys. Res. 115, C06023 (2010).ADS 

Google Scholar 
Iacono, R., Napolitano, E., Marullo, S., Artale, V. & Vetrano, A. Seasonal variability of the Tyrrhenian Sea surface geostrophic circulation as assessed by altimeter data. J. Phys. Oceanogr. 43, 1710–1732. https://doi.org/10.1175/JPO-D-12-0112.1 (2013).ADS 
Article 

Google Scholar 
Boero, F. et al. CoCoNet: Towards coast to coast networks of Marine Protected Areas (from the shore to the high and deep sea), coupled with sea-based wind energy potential. Sci. Res. Inf. Technol. 6(Suppl.), 1–95 (2016).
Google Scholar 
Rio, M. H. et al. A mean dynamic topography of the Mediterranean Sea computed from altimetric data, in-situ measurements and a general circulation model. J. Mar. Syst. 65, 484–508 (2007).Article 

Google Scholar 
Cucco, A. et al. Hydrodynamic modelling of coastal seas: The role of tidal dynamics in the Messina Strait, Western Mediterranean Sea. Nat. Hazards Earth Syst. Sci. 16, 1553–1569 (2016).ADS 
Article 

Google Scholar 
Hopkins, T. S., Salusti, E. & Settimi, D. Tidal forcing of the water mass interface in the Straits of Messina. J. Geophys. Res. 89, 2013–2024 (1984).ADS 
Article 

Google Scholar 
Bignami, F. & Salusti, E. Tidal currents and transient phenomena in the Strait of Messina: A review. In: The Physical Oceanography of Sea Straits (ed. Pratt, L. J.) 95–124 (Kluwer Academic, 1990).Azzaro, F., Decembrini, F., Raffa, F. & Crisafi, E. Seasonal variability of phytoplankton fluorescence in relation to the Straits of Messina (Sicily) tidal upwelling. Ocean Sci. Discuss. 4, 415–440 (2007).ADS 

Google Scholar 
De Domenico, E., Cortese, G. & Pulicanò, G. Chemical characteristics of the waters in the Straits of Messina. In The Straits of Messina ecosystem, present knowledge for an ecohydrodynamical approach. Proceedings of Symposium held in Messina, 4–6 April 1991, Messina (eds. Guglielmo, L., Manganaro, A., & De Domenico, E.) 155–167 (Dipartimento di Biologia Animale ed Ecologia Marina, 1995).Guglielmo, L. Distribuzione di Chetognati nell’area idrografica dello Stretto di Messina. Pubbl. Staz. Zool. Napoli 40, 34–72 (1976).
Google Scholar 
Sitran, R., Bergamasco, A., Decembrini, F. & Guglielmo, L. Temporal succession of tintinnids in the northern Ionian Sea, Central Mediterranean. J. Plankton Res. 29, 495–508 (2007).Article 

Google Scholar 
AA.VV. Final Scientific Report of the Project Cluster 10—SAM “Realizzazione ed attivazione di una rete integrata di piattaforme costiere e mezzo mobile attrezzati per Sistemi Avanzati di Monitoraggio delle acque (SAM)”, funded by the Italian Ministry of University and Scientifical and Technological Research (MURST), Internal Data File, Istituto Sperimentale Talassografico, National Research Council, Messina, Italy (2005).Sitran, R. Caratterizzazione dei popolamenti microzooplanctonici nell’area idrografica dello Stretto di Messina, University of Messina, Ph.D. Thesis XVII cycle (2006) (in Italian).Bergamasco, A. et al. A laboratory for the observation of a highly-energetic coastal marine system: The Straits of Messina. In Volume DTA/06–2011, “Marine Research at CNR” 2185–2202 (Department of Earth and Environment of National Research Council, 2011).Doyle, T. K. et al. Widespread occurrence of the jellyfish Pelagia noctiluca in Irish coastal and shelf waters. J. Plankton Res. 30, 963–968 (2008).Article 

Google Scholar 
Guglielmo, L. Spiaggiamenti di eufausiacei lungo la costa messinese dello Stretto dal dicembre 1968 al dicembre 1969. Boll. Pesca Piscic. Idrobiol. 24, 71–77 (1969).
Google Scholar 
Guglielmo, L., Costanzo, G. & Berdar, A. Ulteriore contributo alla conoscenza dei crostacei spiaggiati lungo il litorale messinese dello Stretto. Atti Soc. Pelorit. 19, 129–156 (1973).
Google Scholar 
Scotto Di Carlo, B., Costanzo, G., Fresi, E., Guglielmo, L. & Ianora, A. Feeding ecology and stranding mechanisms in two lanternfishes, Hygophum benoiti and Myctophum punctatum. Mar. Ecol. Prog. Ser 9, 13–24 (1982).ADS 
Article 

Google Scholar 
Battaglia, P., Ammendolia, G., Cavallaro, M., Consoli, P. & Esposito, V. Influence of lunar phases, winds and seasonality on the stranding of mesopelagic fish in the Strait of Messina (Central Mediterranean Sea). Mar. Ecol. 38, e12459. https://doi.org/10.1111/maec.12459 (2017).Article 

Google Scholar 
Umgiesser, G., Canu, D. M., Cucco, A. & Solidoro, C. A finite element model for the Venice Lagoon. Development, set up, calibration and validation. J. Mar. Syst. 51, 123–145 (2004).Article 

Google Scholar 
Ferrarin, C., Bergamasco, A., Umgiesser, G. & Cucco, A. Hydrodynamics and spatial zonation of the Capo Peloro coastal system (Sicily) through 3-D numerical modeling. J. Mar. Syst. 117, 96–107 (2013).Article 

Google Scholar 
Umgiesser, G., Ferrarin, C., Cucco, A., De Pascalis, F. & Bellafiore, D. Comparative hydrodynamics of 10 Mediterranean lagoons by means of numerical modeling. J. Geophys. Res. Oceans 119, 2212–2226 (2014).ADS 
Article 

Google Scholar 
Cucco, A., Quattrocchi, G., Satta, A., Antognarelli, F. & De Biasio, F. Predictability of wind-induced sea surface transport in coastal areas. J. Geophys. Res. Oceans 121, 5847–5871. https://doi.org/10.1002/2016JC011643 (2016).ADS 
Article 

Google Scholar 
Cucco, A., Quattrocchi, G. & Zecchetto, S. The role of temporal resolution in modeling the wind induced sea surface transport in coastal seas. J. Mar. Syst. 193, 46–58. https://doi.org/10.1016/j.jmarsys.2019.01.004 (2019).Article 

Google Scholar 
Quattrocchi, G. et al. An operational numerical system for oil stranding risk assessment in a high-density vessel traffic area. Front. Mar. Sci. 8, 585396. https://doi.org/10.3389/fmars.2021.585396 (2021).Article 

Google Scholar 
Cucco, A. et al. A high-resolution real-time forecasting system for predicting the fate of oil spills in the Strait of Bonifacio (western Mediterranean Sea). Mar. Pollut. Bull. 64, 1186–1200 (2012).CAS 
PubMed 
Article 

Google Scholar 
Cucco, A. & Umgiesser, G. The Trapping Index: How to integrate the Eulerian and the Lagrangian approach for the computation of the transport time scales of semi-enclosed basins. Mar. Pollut. Bull. 98, 210–220 (2015).CAS 
PubMed 
Article 

Google Scholar 
Quattrocchi, G. et al. Optimal design of a Lagrangian observing system for hydrodynamic surveys. J. Oper. Oceanogr. 9(suppl.), s77–s88. https://doi.org/10.1080/1755876X.2015.1114805 (2016).Article 

Google Scholar 
Quattrocchi, G. et al. Hydrodynamic controls on connectivity of the high commercial value shrimp Parapenaeus longirostris (Lucas, 1846) in the Mediterranean Sea. Sci. Rep. 9, 16935. https://doi.org/10.1038/s41598-019-53245-8 (2019).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pastor-Prieto, M. et al. Spatial heterogeneity of Pelagia noctiluca ephyrae linked to water masses in the Western Mediterranean. PLoS ONE 16, e0249756. https://doi.org/10.1371/journal.pone.0249756 (2021).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Haeckel, E. Das system der medusen. Monographie der Medusen 499–510 (Gustav Fischer Verlag, 1880).
Google Scholar 
Avian, M. Temperature influence on in vitro reproduction and development of Pelagia noctiluca (Forsskäl, 1775). Boll. Zool. 53, 385–391 (1986).Article 

Google Scholar 
Fossette, S. et al. Current-oriented swimming by jellyfish and its role in bloom maintenance. Curr. Biol. 25, 342–347. https://doi.org/10.1016/j.cub.2014.11.050 (2015).CAS 
Article 
PubMed 

Google Scholar 
Pinardi, N. et al. Mediterranean Sea large-scale low-frequency ocean variability and water mass formation rates from 1987 to 2007: A retrospective analysis. Prog. Oceanogr. 132, 318–332 (2015).ADS 
Article 

Google Scholar 
Demirov, E. & Pinardi, N. Simulation of the Mediterranean Sea circulation from 1979 to 1993: Part I. The interannual variability. J. Mar. Syst. 33–34, 23–50 (2002).Article 

Google Scholar 
Menna, M. et al. New insights of the Sicily channel and Southern Tyrrhenian sea variability. Water 11, 1355 (2019).Article 

Google Scholar 
Avian, M. & Rottini Sandrini, L. Oocyte development in four species of scyphomedusa in the northern Adriatic Sea. Hydrobiologia 216/217, 189–195 (1991).Article 

Google Scholar 
Malej, A. Behaviour and trophic ecology of the jellyfish Pelagia noctiluca (Forsskäl, 1775). J. Exp. Mar. Biol. Ecol. 126, 259–270 (1989).Article 

Google Scholar 
Lo Bianco, S. Notizie biologiche riguardanti specialmente il periodo di maturità sessuale degli animali del golfo di Napoli. Mitt. Zool. Stn. Neapel 19, 513–761 (1909).
Google Scholar 
Purcell, J. E., Malej, A. & Benović, A. Potential links of jellyfish to eutrophication and fisheries. In Coastal and Estuarine Studies, Ecosystem at the Land-Sea Margin Drainage Basin to Coastal Sea (eds Malone, T. C. et al.) 241–263 (American Geophysical Union, 1999).Chapter 

Google Scholar 
Spezie, G. C., Sansone, E., Budillon, G. & Gallarato, A. Caratterizzazione idrodinamica del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia. Campagna oceanografica 1994. Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM94). In Data Rep., (eds. Faranda, F. M. & Povero, P.) 1–82 (1995).Spezie, G. C. et al. Rilievi idrodinamici nel sistema Eolie e nei bacini limitrofi di Cefalù e Gioia. Campagna oceanografiche 1995. Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 1–98 (1996).Carrada, G. C., Ribera D’Alcalà, M. & Saggiomo, V. The pelagic system of the Southern Tyrrhenian Sea. Some comments and working hypotheses. In Proceedings IX Proceedings XII Italian Association of Oceanography and Limnology Congress 151–166 (1992).Povero, P., Misic, C., Acconero, A. & Fabiano M. Distribuzione e caratterizzazione biochimica della sostanza organica particellata nelle acque del Tirreno Sud Orientale. In Acts 12 Congress of the Italian Association of Oceanology and Limnology 227–237 (1998).Brancato, G., Minutoli, R., Granata, A., Sidoti, O. & Guglielmo L. Diversity and vertical migration of euphausiids across the Straits of Messina area. In: Mediterranean Ecosystem: Structures and Processes (eds. Faranda, F. M., Guglielmo, L. & Spezie, G.) 131–141 (Springer, 2001).Sitran, R., Bergamasco, A., Decembrini, F. & Guglielmo, L. Microzooplankton (tintinnid ciliates) diversity: Coastal community structure and driving mechanisms in the Southern Tyrrhenian Sea (Western Mediterranean). J. Plankton Res. 31, 153–170 (2009).Article 

Google Scholar 
Fonda Umani, S., Monti, M., Minutoli, R. & Guglielmo, L. Recent advances in the Mediterranean researches on zooplankton: from spatial–temporal patterns of distribution to processes oriented studies. Adv. Oceanogr. Limnol. 1, 295–356 (2010).Article 

Google Scholar 
Giordano, D. et al. Summer larval fish assemblages in the Southern Tyrrhenian Sea (Western Mediterranean Sea). Mar. Ecol. 36, 104–117. https://doi.org/10.1111/maec.12123 (2015).ADS 
Article 

Google Scholar 
Fonda Umani, S., Milani, L. & Martecchini, E. Distribuzione dei popolamenti microzooplanctonici durante la campagna oceanografica Eolie 1994. Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 199–222 (1995).Carrada, G. C., Mangoni, O. & Sgrosso, S. Distribuzione spaziale di clorofilla a e di feopigmenti in diverse frazioni dimensionali del fitoplancton. Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 197–216 (1996).Guglielmo, L. et al. Distribuzione verticale e migrazione giornaliera dello zooplancton e del micronecton nel Tirreno meridionale (Isole Eolie). Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 217–246 (1996).Innamorati, M., Lazzara, L., Massi, L., Biondi, N. & Nuccio, C. Fitoplancton, luce e produzione primaria nella’Arcipelago delle Isole Eolie, in estate. Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 161–196 (1996).Zunini Sertorio, T., Licandro, P., Giallain, M. & Bernat, P. Distribuzione verticale della biomassa zooplanctonica su una stazione delle Isole Eolie (Luglio 1995). Caratterizzazione ambientale marina del sistema Eolie e dei bacini limitrofi di Cefalù e Gioia (EUCUMM95). In Data Rep. (eds. Faranda, F. M. & Povero, P.) 247–254 (1996).Sabates, A. et al. Pathways for Pelagia noctiluca jellyfish intrusions onto the Catalan shelf and their interactions with early life fish stages. J. Mar. Syst. 187, 52–61 (2018).Article 

Google Scholar 
Mosetti, F. Currents in the Straits of Messina. In The Straits of Messina ecosystem (eds Guglielmo, L. et al.) 13–29 (University of Messina, Department of Marine Biology and Ecology, 1995).
Google Scholar 
Zavodnik, D. Spatial aggregations of the swarming jellyfish Pelagia noctiluca (Scyphozoa). Mar. Biol. 94, 265–269 (1987).Article 

Google Scholar 
El Rahi, J., Weeber, M. P. & El Serafy, G. Modelling the effect of behavior on the distribution of the jellyfish Mauve stinger (Pelagia noctiluca) in the Balearic Sea using an individual-based model. Ecol. Model. 433, 109230 (2020).Article 

Google Scholar 
Axiak, V. & Civili, F. S. Jellyfish blooms in the Mediterranean: causes, mechanisms, impact on man and the environment. A programme review. In: UNEP: Jellyfish blooms in the Mediterranean. Proceedings of the II Workshop on Jellyfish in the Mediterranean Sea. MAP Tech. Rep. Ser. Vol. 47, 1–10 (UNEP, 1991).Keesing, J. K. et al. Role of winds and tides in timing of beach strandings, occurrence, and significance of swarms of the jellyfish Crambione mastigophora Mass 1903 (Scyphozoa: Rhizostomeae: Catostylidae) in north-western Australia. Hydrobiologia 768, 19–36. https://doi.org/10.1007/s10750-015-2525-5 (2016).CAS 
Article 

Google Scholar 
Aglieri, G. et al. First evidence of inbreeding, relatedness and chaotic genetic patchiness in the holoplanktonic jellyfish Pelagia noctiluca (Scyphozoa, Cnidaria). PLoS ONE 9, e99647. https://doi.org/10.1371/journal.pone.0099647 (2014).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Alpers, W., Brandt, P. & Rubino, A. Internal waves generated in the Strait of Gibraltar and Messina: Observations from space. In Remote Sensing of the European Seas (eds. Barale, V. & Gade, M.) 319–330 (Springer, 2008). https://doi.org/10.1007/978-1-4020-6772.Droghei, R. et al. The role of Internal Solitary Waves on deep-water sedimentary processes: The case of up-slope migrating sediment waves off the Messina Strait. Sci. Rep. 6, 36376. https://doi.org/10.1038/srep36376 (2016).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
La Forgia, G. et al. Sediment resuspension and bedform generation induced by internal solitary waves. Geophys. Res. Abs. Vol. 21, EGU2019-9121, EGU General Assembly (2019).Lohmann, H. Die Stromunger in der Strasse von Messina und die verteilung des planktons in derselben. Int. Rev. Ges. Hydrobiol. 2, 505–556 (1909).Article 

Google Scholar 
Magazzù, G. & Andreoli, C. Trasferimenti fitoplanctonici attraverso lo Stretto di Messina in relazione alle condizioni idrologiche. Boll. Pesca Piscic. Idrobiol. 26, 125–193 (1971).
Google Scholar 
Palanques, A. et al. General patterns of circulation, sediment fluxes and ecology of the Palamòs (La Fonera) submarine canyon, northwestern Mediterranean. Progr. Oceanogr. 66, 89–119 (2005).ADS 
Article 

Google Scholar 
Granata, A. et al. Vertical distribution and diel migration of zooplankton and micronekton in Polcevera submarine canyon of the Ligurian mesopelagic zone (NW Mediterranean Sea). Progr. Oceanogr. 183, 102298. https://doi.org/10.1016/j.pocean (2020).Article 

Google Scholar 
Zagami, G. et al. Spring copepod vertical zonation pattern and diel migration in the open Ligurian Sea (north-western Mediterranean). Progr. Oceanogr. 183, 102297. https://doi.org/10.1016/j.pocean (2020).Article 

Google Scholar 
Danovaro, R. & Boero, F. Italian seas. In: World Seas: An Environmental Evaluation. Vol. I Europe, The Americas and West Africa. (ed. Sheppard, C.) 283–306 (Elsevier Ltd., 2019). https://doi.org/10.1016/B978-0-12-805068-2.00044-9Lo Iacono, C., Sulli, A. & Agate, M. Submarine canyons of north-western Sicily (Southern Tyrrhenian Sea): Variability in morphology, sedimentary processes and evolution on a tectonically active margin. Deep-Sea Res. 104, 93–105 (2014).
Google Scholar  More

Försterra, G. et al. Animal forests in the Chilean fiord region: Discoveries and perspectives in shallow and deep waters. In Marine Animal Forests. Orejas Saco del Valle (eds Rossi, S. et al.) 1–35 (Springer, 2016). https://doi.org/10.1007/978-3-319-17001-5_3-1.Chapter 

Google Scholar 
Castilla, J. C. et al. (eds) Conservación en la Patagonia Chilena: Evaluación del conocimiento, oportunidades y desafíos (Ediciones Universidad Católica, 2021).
Google Scholar 
Iriarte, J. L. et al. Oceanographic Processes in Chilean Fjords of Patagonia: From small to large-scale studies. Prog. Oceanogr. 129, 1–7. https://doi.org/10.1016/j.pocean.2014.10.004 (2014).ADS 
Article 

Google Scholar 
Iriarte, J. L. Natural and human influences on marine processes in Patagonian Subantarctic coastal waters. Front. Mar. Sci. 5, 360. https://doi.org/10.3389/fmars.2018.00360 (2018).Article 

Google Scholar 
Strub, P. T. et al. Ocean circulation along the southern Chile transition region (38°–46°S): Mean, seasonal and interannual variability, with a focus on 2014–2016. Prog. Oceanogr. 172, 159–198. https://doi.org/10.1016/j.pocean.2019.01.004 (2019).ADS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Häussermann, V. et al. Macrobentos de fondos duros de la Patagonia chilena: Énfasis en la conservación de bosques sublitorales de invertebrados y algas. In Conservación en la Patagonia Chilena: Evaluación del conocimiento, oportunidades y desafíos (eds Castilla, J. C. et al.) (Ediciones Universidad Católica, 2021).
Google Scholar 
Kol, P. H. Los Riesgos de la Expansión Salmonera en la Patagonia Chilena. Estado de la Salmonicultura Intensiva en la Región de Magallanes (AIDA, 2018).Iversen, A. et al. Production cost and competitiveness in major salmon farming countries 2003–2018. Aquaculture 522, 735089. https://doi.org/10.1016/j.aquaculture.2020.735089 (2020).Article 

Google Scholar 
Cárdenas-Retamal, R. et al. Impact assessment of salmon farming on income distribution in remote coastal areas: The Chilean case. Food Policy 101, 102078. https://doi.org/10.1016/j.foodpol.2021.102078 (2021).Article 

Google Scholar 
Chavez, C. et al. Main issues and challenges for sustainable development of salmon farming in Chile: A socio-economic perspective. Rev. Aquac. 11, 403–421. https://doi.org/10.1111/raq.12338 (2019).Article 

Google Scholar 
Quiñones, R. A. et al. Environmental issues in Chilean salmon farming: A review. Rev. Aquac. 11, 375–402. https://doi.org/10.1111/raq.12337 (2019).Article 

Google Scholar 
Mardones, J. I. et al. Disentangling the environmental processes responsible for the world’s largest farmed fish-killing harmful algal bloom: Chile, 2016. Sci. Total Environ. 76, 1–19. https://doi.org/10.1016/j.scitotenv.2020.144383 (2021).CAS 
Article 

Google Scholar 
Navedo, J. G. et al. Upraising a silent pollution: Antibiotic resistance at coastal environments and transference to long-distance migratory shorebirds. Sci. Total Environ. 777, 1–7. https://doi.org/10.1016/j.scitotenv.2021.146004 (2021).CAS 
Article 

Google Scholar 
SUBPESCA. Listado de concesiones de acuicultura d salmónidos por agrupación de concesiones en las regiones X, XI y XII (Julio 2021). https://www.subpesca.cl/portal/619/w3-article-103129.html (2021).Gorny, M. et al. Las comunidades marinas bentónicas de la Reserva Nacional Katalalixar (Chile). Oceanografía, 29–44 (2020).Friedlander, A. M. et al. Marine communities of the newly created Kawésqar National Reserve, Chile: From glaciers to the Pacific Ocean. PLoS One 16(4), e0249413. https://doi.org/10.1371/journal.pone.0249413 (2021).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Mardones, J. I. et al. Toxic dinoflagellate blooms of Alexandrium catenella in Chilean fjords: A resilient winner from climate change. ICES J. Mar. Sci. 74(4), 988–995. https://doi.org/10.1093/icesjms/fsw164 (2016).Article 

Google Scholar 
Alvarez-Garreton, C. et al. The CAMELS-CL dataset: Catchment attributes and meteorology for large sample studies—Chile dataset. Hydrol. Earth Syst. Sci. 22, 5817–5846. https://doi.org/10.5194/hess-22-5817-2018 (2018).ADS 
Article 

Google Scholar 
Novak, B. J. et al. Transforming ocean conservation: Applying the genetic rescue toolkit. Genes 11, 209. https://doi.org/10.3390/genes11020209 (2020).CAS 
Article 
PubMed Central 

Google Scholar 
Outeiro, L. et al. Using ecosystem services mapping for marine spatial planning in southern Chile under scenario assessment. Ecosyst. Serv. 16, 341–353. https://doi.org/10.1016/j.ecoser.2015.03.004 (2015).Article 

Google Scholar 
Anbleyth-Evans, J. et al. Toward marine democracy in Chile: Examining aquaculture ecological impacts through common property local ecological knowledge. Mar. Policy 113, 103690. https://doi.org/10.1016/j.marpol.2019.103690 (2019).Article 

Google Scholar 
Kershaw, F. et al. Geospatial genetics: Integrating genetics into marine protection and spatial planning. Aquat. Conserv. Mar Freshw. Ecosyst. https://doi.org/10.1002/aqc.3622 (2021).Article 

Google Scholar 
Jenkins, T. L. & Stevens, J. R. Assessing connectivity between MPAs: Selecting taxa and translating genetic data to inform policy. Mar. Policy 94, 165–173. https://doi.org/10.1016/j.marpol.2018.04.022 (2018).Article 

Google Scholar 
Paredes, J. et al. Population genetic structure at the northern edge of the distribution of Alexandrium catenella in the Patagonian fjords and its expansion along the open Pacific Ocean coast. Front. Mar. Sci. 5, 532. https://doi.org/10.3389/fmars.2018.00532 (2019).Article 

Google Scholar 
Canales-Aguirre, C. B. C. et al. Population genetic structure of Patagonian toothfish (Dissostichus eleginoides) in the Southeast Pacific and Southwest Atlantic Ocean. PeerJ 6, e4173. https://doi.org/10.7717/peerj.4173 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Canales-Aguirre, C. B. C. et al. High genetic diversity and low-population differentiation in the Patagonian sprat (Sprattus fuegensis) based on mitochondrial DNA. Mitochondrial DNA Part A 29(8), 1148–1155. https://doi.org/10.1080/24701394.2018.1424841 (2018).CAS 
Article 

Google Scholar 
Pérez-Alvarez, M. et al. Historical dimensions of population structure in a continuously distributed marine species: The case of the endemic Chilean dolphin. Sci. Rep. 6, 35507. https://doi.org/10.1038/srep35507 (2016).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pérez-Alvarez, J. M. et al. Phylogeography and demographic inference of the endangered sei whale, with implications for conservation. Aquat. Conserv. Mar. Freshw. Ecosyst. https://doi.org/10.1002/aqc.3717 (2021).Article 

Google Scholar 
Addamo, A. M. et al. Global-scale genetic structure of a cosmopolitan cold-water coral species. Aquat. Conserv. Mar. Freshw. Ecosyst. 31(1), 1–14. https://doi.org/10.1002/aqc.3421 (2021).Article 

Google Scholar 
Addamo, A. M. et al. Genetic conservation management of marine resources and ecosystems of Patagonian Fjords. Front. Mar. Sci. 8, 612195. https://doi.org/10.3389/fmars.2021.612195 (2021).Article 

Google Scholar 
Addamo, A. M. et al. Development of microsatellite markers in the deep-sea cup coral Desmophyllum dianthus and cross-species amplifications in the Scleractinia Order. J. Hered. 106(3), 322–330. https://doi.org/10.1093/jhered/esv010 (2015).CAS 
Article 
PubMed 

Google Scholar 
Miller, K. J. & Gunasekera, R. M. A comparison of genetic connectivity in two deep sea corals to examine whether seamounts are isolated islands or stepping stones for dispersal. Sci. Rep. 7, 1–14. https://doi.org/10.1038/srep46103 (2017).CAS 
Article 

Google Scholar 
Holloley, C. E. & Geerts, P. G. Multiplex Manager 1.0: A cross-platform computer program that plans and optimizes multiplex PCR. Biotechniques 46, 511–517. https://doi.org/10.2144/000113156 (2009).Article 

Google Scholar 
Brookfield, J. F. Y. A simple new method for estimating null allele frequency from heterozygote deficiency. Mol. Ecol. 5, 453–455. https://doi.org/10.1046/j.1365-294X.1996.00098.x (1996).CAS 
Article 
PubMed 

Google Scholar 
Van Oosterhout, C. et al. Micro-Checker: Software for identifying and correcting genotyping errors in microsatellite data. Mol. Ecol. Notes 4, 535–538. https://doi.org/10.1111/j.1471-8286.2004.00684.x (2004).CAS 
Article 

Google Scholar 
Chapuis, M.-P. & Estoup, A. Microsatellite null alleles and estimation of population differentiation. Mol. Biol. Evol. 24(3), 621–631 (2007).CAS 
Article 

Google Scholar 
Peakall, R. & Smouse, P. E. GenAlEx 6.5: Genetic analysis in Excel. Population genetic software for teaching and research—An update. Bioinformatics 28, 2537–2539. https://doi.org/10.1093/bioinformatics/bts460 (2012).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Rousset, F. Genepop’007: A complete re-implementation of the Genepop software for Windows and Linux. Mol. Ecol. Resour. 8, 103–106. https://doi.org/10.1111/j.1471-8286.2007.01931.x (2008).Article 
PubMed 

Google Scholar 
Holm, S. A simple sequentially rejective multiple test procedure. Scand. J. Stat. 6, 65–70 (1979).MathSciNet 
MATH 

Google Scholar 
Excoffier, L. & Lischer, H. E. L. Arlequin suite ver 3.5: A new series of programs to perform population genetics analyses under Linux and Windows. Mol. Ecol. Resour. 10, 564–567. https://doi.org/10.1111/j.1755-0998.2010.02847.x (2010).Article 
PubMed 

Google Scholar 
Falush, D. et al. Inference of population structure using multilocus genotype data: Linked loci and correlated allele frequencies. Genetics 164, 1567–1587. https://doi.org/10.1111/j.1471-8286.2007.01758.x (2003).CAS 
Article 
PubMed 
PubMed Central 

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, 359–361. https://doi.org/10.1007/s12686-011-9548-7 (2012).Article 

Google Scholar 
Li, Y. L. & Liu, J. X. StructureSelector: A web based software to select and visualize the optimal number of clusters using multiple methods. Mol. Ecol. Resour. 18, 176–177. https://doi.org/10.1111/1755-0998.12719 (2018).Article 
PubMed 

Google Scholar 
Kopelman, N. M. et al. CLUMPAK: A program for identifying clustering modes and packaging population structure inferences across K. Mol. Ecol. Resour. 15, 1179–1191. https://doi.org/10.1111/1755-0998.12387 (2015).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pritchard, J. K. et al. Inference of population structure using multilocus genotype data. Genetics 155, 945–959 (2000).CAS 
Article 

Google Scholar 
Evanno, G. et al. Detecting the number of clusters of individuals using the software STRUCTURE: A simulation study. Mol. Ecol. 14, 2611–2620. https://doi.org/10.1111/j.1365-294X.2005.02553.x (2005).CAS 
Article 
PubMed 

Google Scholar 
Puechmaille, S. J. The program structure does not reliably recover the correct population structure when sampling is uneven: Subsampling and new estimators alleviate the problem. Mol. Ecol. Resour. 16, 608–627. https://doi.org/10.1111/1755-0998 (2016).Article 
PubMed 

Google Scholar 
Piry, S. et al. GeneClass2: A software for genetic assignment and first-generation migrant detection. J. Hered. 95, 536–539. https://doi.org/10.1093/jhered/esh074 (2004).CAS 
Article 
PubMed 

Google Scholar 
Cornuet, J. M. & Luikart, G. Description and power analysis of two tests for detecting recent population bottlenecks from allele frequency data. Genetics 144, 2001–2014 (1997).Article 

Google Scholar 
Wickham, H. et al. Welcome to the tidyverse. J. Open Source Softw. 4(43), 1686. https://doi.org/10.21105/joss.01686 (2019).ADS 
Article 

Google Scholar 
Wickham, H. et al. dplyr: A grammar of data manipulation. https://dplyr.tidyverse.org, https://github.com/tidyverse/dplyr (2022).Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer, 2022). https://ggplot2.tidyverse.org. ISBN 978-3-319-24277-4.Addamo, A. M. et al. Microsatellites of Desmophyllum dianthus—Comau Fjord. Front. Mar. Sci. https://doi.org/10.3389/fmars.2021.612195. Zenodo. https://doi.org/10.5281/zenodo.4435966 (2021).Tecklin, D. Sensing the limits of fixed marine property rights in changing coastal ecosystems: Salmon aquaculture concessions, crises, and governance challenges in Southern Chile. J. Int. Wildl. Law Policy 19(4), 284–300. https://doi.org/10.1080/13880292.2016.1248647 (2016).Article 

Google Scholar 
Buschmann, A. H. et al. Salmon aquaculture and coastal ecosystem health in Chile: Analysis of regulations, environmental impacts and bioremediation systems. Ocean Coast. Manag. 52, 243–249. https://doi.org/10.1016/j.ocecoaman.2009.03.002 (2009).Article 

Google Scholar 
Pantoja, S. et al. Oceanography of the Chilean Patagonia. Cont. Shelf Res. 31, 149–153. https://doi.org/10.1016/j.csr.2010.10.013 (2011).ADS 
Article 

Google Scholar 
Molina, V. & Fernández, C. Bacterioplankton response to nitrogen and dissolved organic matter produced from salmon mucus. Microbiol. Open 9(12), e1132. https://doi.org/10.1002/mbo3.1132 (2020).CAS 
Article 

Google Scholar 
Försterra, G. & Häussermann, V. First report on large scleractinian (Cnidaria: Anthozoa) accumulations in cold-temperate shallow water of south Chilean fjords. Zool. Verh. 345, 117–128 (2003).
Google Scholar 
Brown, S. M. et al. Limited population structure, genetic drift and bottlenecks characterise an endangered bird species in a dynamic, fire-prone ecosystem. PLoS One 8(4), e59732. https://doi.org/10.1371/journal.pone.0059732 (2013).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Takahashi, Y. et al. Lack of genetic variation prevents adaptation at the geographic range margin in a damselfly. Mol. Ecol. 25, 4450–4460. https://doi.org/10.1111/mec.13782 (2016).Article 
PubMed 

Google Scholar 
Thiel, M. et al. The Humboldt Current system of Northern and Central Chile. Oceanographic processes, ecological interactions and socioeconomic feedback. Oceanogr. Mar. Biol. Annu. Rev. 45, 195–344 (2007).
Google Scholar 
Giesecke, R. et al. General Hydrography of the Beagle Channel, a Subantarctic Interoceanic Passage at the Southern Tip of South America. Front. Mar. Sci. Coast. Ocean Process. 8, 621822. https://doi.org/10.3389/fmars.2021.621822 (2021).Article 

Google Scholar 
Chaigneau, A. Surface circulation and fronts of the South Pacific Ocean, east of 120°. Geophys. Res. Lett. 32, L08605. https://doi.org/10.1029/2004GL022070 (2005).ADS 
Article 

Google Scholar 
Aiken, C. M. A reanalysis of the Chilean ocean circulation: Preliminary results for the region between 20°S to 40°S. Lat. Am. J. Aquat. Res. 45(1), 193–198. https://doi.org/10.3856/vol45-issue1-fulltext-19 (2017).Article 

Google Scholar 
González, H. E. et al. Primary production and plankton dynamics in the Reloncaví Fjord and the Interior Sea of Chiloé, Northern Patagonia, Chile. Mar. Ecol. Prog. Ser. 402, 13–30. https://doi.org/10.3354/meps08360 (2010).ADS 
CAS 
Article 

Google Scholar 
González, H. E. et al. Seasonal plankton variability in Chilean Patagonia fjords: Carbon flow through the pelagic food web of Aysen Fjord and plankton dynamics in the Moraleda Channel basin. Cont. Shelf Res. 31, 225–243. https://doi.org/10.1016/j.csr.2010.08.010 (2011).ADS 
Article 

Google Scholar 
Feehan, K. A. et al. Highly seasonal reproduction in deep-water emergent Desmophyllum dianthus (Scleractinia: Caryophylliidae) from the Northern Patagonian Fjords. Mar. Biol. 166(4), 52. https://doi.org/10.1007/s00227-019-3495-3 (2019).Article 

Google Scholar 
Försterra, G. et al. Mass die off of the cold-water coral Desmophyllum dianthus in the Chilean Patagonian Fjord Region. Bull. Mar. Sci. 90(3), 895–899 (2014).Article 

Google Scholar 
Mora-Soto, A. et al. A song of wind and ice: Increased frequency of marine cold-spells in southwestern Patagonia and their possible effects on giant kelp forests. J. Geophys. Res. Oceans 127, e2021JC017801. https://doi.org/10.1029/2021JC017801 (2022).ADS 
Article 

Google Scholar 
Brown, J. H. On the relationship between abundance and distribution of species. Am. Nat. 124, 255–279 (1984).Article 

Google Scholar 
Verberk, W. Explaining general patterns in species abundance and distributions. Nat. Sci. Educ. 3(10), 38 (2011).
Google Scholar 
Devenish, C. et al. Extreme and complex variation in range-wide abundances across a threatened Neotropical bird community. Divers. Distrib. 23, 910–921. https://doi.org/10.1111/ddi.12577 (2017).Article 

Google Scholar 
Iriarte, J. L. et al. Influence of seasonal freshwater streamflow regimes on phytoplankton blooms in a Patagonian fjord. N. Z. J. Mar. Freshw. Res. 51(2), 304–315. https://doi.org/10.1080/00288330.2016.1220955 (2016).CAS 
Article 

Google Scholar 
Silva, N. et al. Características oceanográficas físicas y químicas de canales australes chilenos entre Puerto Montt y Laguna San Rafael (Crucero Cimar-Fiordo 1). Cienc. Tecnol. Mar. 20, 23–106 (1997).
Google Scholar 
Iriarte, J. L. et al. Low spring primary production and microplankton carbon biomass in Sub-Antarctic Patagonian channels and fjords (50–53°S). Arct. Antarct. Alp. Res. 50(1), e1525186. https://doi.org/10.1080/15230430.2018.1525186 (2018).Article 

Google Scholar 
Höfer, J. et al. All you can eat: The functional response of the cold-water coral Desmophyllum dianthus feeding on krill and copepods. PeerJ 6, e5872. https://doi.org/10.7717/peerj.5872 (2018).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Montero, P. et al. A winter dinoflagellate bloom drives high rates of primary production in a Patagonian fjord ecosystem. Estuar. Coast. Shelf Sci. 199, 105e116. https://doi.org/10.1016/j.ecss.2017.09.027 (2017).CAS 
Article 

Google Scholar 
Quiroga, E. et al. Seasonal benthic patterns in a glacial Patagonian fjord: The role of suspended sediment and terrestrial organic matter. Mar. Ecol. Prog. Ser. 561, 31–50. https://doi.org/10.3354/meps11903 (2016).ADS 
Article 

Google Scholar 
Escribano, R. et al. Seasonal and inter-annual variation of mesozooplankton in the coastal upwelling zone off central-southern Chile. Prog. Oceanogr. 75, 470–485. https://doi.org/10.1016/j.pocean.2007.08.027 (2007).ADS 
Article 

Google Scholar 
Gori, A. et al. Physiological response of the cold-water coral Desmophyllum dianthus to thermal stress and ocean acidification. PeerJ 4, e1606. https://doi.org/10.7717/peerj.1606 (2016).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Martínez-Dios, A. et al. Effects of low pH and feeding on calcification rates of the cold-water coral Desmophyllum dianthus. PeerJ 8, e8236. https://doi.org/10.7717/peerj.8236 (2020).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
López-Márquez, V. et al. Asexual reproduction in bad times? The case of Cladocora caespitosa in the eastern Mediterranean Sea. Coral Reefs 40, 663–677. https://doi.org/10.1007/s00338-020-02040-3 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Silva, N. & Calvete, C. Características oceanográficas físicas y químicas de canales australes chilenos entre el Golfo de Penas y el Estrecho de Magallanes (Crucero Cimar-Fiordo 2). Cienc. Tecnol. Mar. 20, 23–88 (2002).
Google Scholar 
Häussermann, V. et al. Species that fly at a higher game: Patterns of deep–water emergence along the Chilean coast, including a global review of the phenomenon. Front. Mar. Sci. 8, 688316. https://doi.org/10.3389/fmars.2021.688316 (2021).Article 

Google Scholar 
Fillinger, L. & Richter, C. Vertical and horizontal distribution of Desmophyllum dianthus in Comau Fjord, Chile: A cold-water coral thriving at low pH. PeerJ 1, e194. https://doi.org/10.7717/peerj.194 (2013).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Addamo, A. M. et al. Biodiversity and distribution of corals in Chile. Mar. Biodivers. 52, 33. https://doi.org/10.1007/s12526-022-01271-7 (2022).Article 

Google Scholar 
Figuerola, B. et al. A review and meta-analysis of potential impacts of ocean acidification on marine calcifiers from the Southern Ocean. Front. Mar. Sci. 8, 584445. https://doi.org/10.3389/fmars.2021.584445 (2021).Article 

Google Scholar 
SGS SIGA. 4.15 Pobreza multidimensional y pobreza por ingresos de la Region de los Lagos. Agosto 2018. Subsecreteria de Desarollo Regional y Administrativo, Gobierno de Chile (2018).FAO. The state of world fisheries and aquaculture. http://www.fao.org/3/a-i720e.pdf (2014).Niklitschek, E. J. et al. Southward expansion of the Chilean salmon industry in the Patagonian Fjords: Main environmental challenges. Rev. Aquac. 4, 1–24. https://doi.org/10.1111/raq.1201 (2013).Article 

Google Scholar 
Soto, M. V. et al. Natural hazards and exposure of strategic connectivity in extreme territories. Comau Fjord, North Patagonia, Chile. Rev. Geogr. Norte Grande 73, 57–75 (2019).Article 

Google Scholar 
Montes, R. M. et al. Quantifying harmful algal bloom thresholds for farmed salmon in southern Chile. Harmful Algae 77, 55–65. https://doi.org/10.1016/j.hal.2018.05.004 (2018).Article 
PubMed 

Google Scholar 
Lembeye, G. Harmful algal blooms in the austral Chilean channels and fjords. In Progress in the Oceanographic Knowledge of Chilean Interior Waters, from Puerto Montt to Cape Horn (eds Silva, N. & Palma, S.) 99–103 (Comité Oceanográfico, 2008).
Google Scholar 
Häussermann, V. et al. Largest baleen whale mass mortality during strong El Niño event is likely related to harmful toxic algal bloom. PeerJ 5, e3123. https://doi.org/10.7717/peerj.3123 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
Google IncGoogle Earth. Retrieved from https://www.google.com/earth/versions/#download-pro (2009). More

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

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

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