Ecology

General description of sequencesAfter the quality filtering step, removal of chimeric fragments, and read merging, a total of 3,378,323 reads with 3007 different features was obtained, with an average of 27,244 sequences per individual sample. After quality filtering, none of the samples was excluded from the analysis of microbial communities.Amoxicillin and thiamphenicol treatments influence microbial diversity and the abundance of specific taxaUsing 16S rRNA NGS, the gut microbial community composition of the chicks in each group was characterized at different time points. At phylum level, microbiota composition varied with age rather than with treatment (Supplementary Fig. S1). Proteobacteria were the most abundant phyla at 1 day of age (d.o.a.), Firmicutes became dominant at later stages, while Bacteroidota were highly abundant in caecum samples collected at 46 d.o.a. Similar dynamics were observed also at family level, since Enterobacteriaceae and Clostridiaceae were significantly more abundant at 1 d.o.a. in all groups, Lactobacillaceae, Lachnospiraceae, and Ruminococcaceae seemed to bloom at 8 d.o.a., and Rikenellaceae were the dominant family in the caecum samples collected at 46 d.o.a. (Fig. 1; Supplementary Fig. S2).Figure 1Heatmap representing the microbial community composition at family level. The heatmap was generated in R (version 4.2.1) (https://www.r-project.org/) using package pheatmap (version 1.0.12).Full size imageEarly-age administrationIn both α-diversity indices (Fig. 2A,B), there was a trend towards increasing diversity from early to late time points in all groups; however, the only significant differences were between the group treated with amoxicillin (AMX1) and the other groups on day 21 post treatment (p.t.), and within AMX1 group between day 21 p.t. and the other time points. PERMANOVA showed that the microbial community was significantly different between the group treated with thiamphenicol (THP1) and the other two groups (i.e. AMX1 and control) on day 1 p.t. (p  More

Total catch control regulation does not lead to the recovery of fisheries and the maintenance of community functionTo contain the decline of wild capture fisheries by overfishing, a series of management regulations have been in place in China to mitigate the fishing impacts as much as possible and maintain sustainable stocks. The “zero-growth” policy is one of the most outstanding representatives. The results showed certain achievements after the implementation of the policy. Simulating the status without the “zero-growth” policy, B/Bmsy fell below 0.5 by 2010 and close to zero by 2019, indicating the impossibility for recovery. However, the policy is not enough for fishery recovery and community health, failing to stop the degradation of fishery resources. Under the implementation of the “zero-growth” policy, B/Bmsy was in a healthy state in 1998, fell below 1 for the first time in 2003, and dropped to 0.52 in 2019, accompanying by F/Fmsy as 1.60. If fishing pressure were maintained at the level of 2019 (F = 1.56 Fmsy), the resource would decline to the depletion state by 2030 (B/Bmsy close to zero, F/Fmsy = 3.64, catch = 35 T). Therefore, a great degree of negative production growth as well as the strict implementation is extremely important. A rapid reduction in the catch control under 0.5 Fmsy scenario would expect to achieve a quick recovery with B/Bmsy over 1 in 2025. Nevertheless, a significant reduction in production would lead to the decline of fishery economics, livelihood difficulties for fishermen and a series of derivative social problems28. An alternative of 1.0 Fmsy would be feasible, under which B/Bmsy could rise to 1 by 2030 with a production of 11.64 MT, close to MSY.The “zero growth” policy faces some inherent challenges, at least from the point of view of ensuring the sustainable use of individual species stocks. Attention should be also paid at the catch quota control of individual species. Because the variation of the intrinsic growth rate of different species, the B is dynamic, and the F changes with the change of B. In a constant production, r-strategic species could remain a higher B/Bmsy than 1 even at a large proportion in catch, but K-strategic species did not show the same fortune. The control of total catch volume rather than individual species could not prevent the community structure from becoming fragile, with the exhaustion of high-trophic species and the decrease of mean trophic level.Individual species have different responses to overfishing that highly associated with their biological characteristicsHigh trophic level species can be sensitive to overfishing, and difficult to rebuild stocks after collapseHairtails Trichiurus spp. are the largest contribution group to China marine capture fisheries, at 0.90 MT about 8.3% of the total production in 20202. They are carnivorous and aggressive with a mean trophic level of 4.4, mainly feeding on fishes in the adult stage, and Mysidacea and Euphausiacea in the juvenile stage29,30. The spawning seasons of Trichiurus spp. are mainly from April to June, and from September to November in Chinese waters31.China coastal areas are excellent foraging and spawning grounds for Trichiurus spp, sustaining a large stock size. If the “zero-growth” policy was not implemented since 1999, the resources of Trichiurus spp. would be exhausted by 2027, having no possibility to recovery at 1.0 Fmsy. Although the total fisheries production has been controlled, and the fishing moratorium period partly covered the spawning seasons of Trichiurus spp., their resource continuous declined into a “destroying” state in 2007, due to the time-lag effect of fishing on high trophic level predators characterized by long population doubling time-consuming32. Under intensive fishing pressure, Trichiurus spp. have showed astonishing fisheries-induced adaption33 by reducing the age and size of maturity, which effectively alleviates the decline rate of B value, resulting the maintenance of Trichiurus spp. capture production. Under the rebuilding scenario of fishing pressure as 1.0 Fmsy, Trichiurus spp. B/Bmsy rose to 0.87 by 2030, lower than the recovery rate of national total capture fisheries, suggesting the recovery rate of high trophic level species could be slow34. Furthermore, in this study fisheries rebuilding only considers the responses of species to fishing pressure, irrespective of a series of factors sensitive to high trophic level species such as pollution and climate change, which indicated a longer period is needed for resource recovery.Middle trophic level species seems non sensitive to total catch control policyAs a representative of middle trophic level species, L. polyactis performed different from Trichiurus spp. Under high fishing pressure. It forms spawning and over-wintering aggregations between nearshore and offshore waters, as well as vertical migration, rising at dusk and falling at dawn35. The spawning season is from mid-February to early May, prior to the national fishing moratorium, indicating young juveniles are in effective protection rather than spawning stock. In the 1950s, L. polyactis was one of the few important species in domestic marine capture fisheries in Chinese waters, producing more than 100,000 T annually5. The catch volumes then showed a downward trend and fell significantly to less than 50,000 T in the 1960–1980s. After 3 decades low catch volumes, the annual capture production rebounded significantly to more than 200,000 T and maintained at such high levels for 2 decades5, showing high resilience to overfishing.Despite many concerns on the risk of resource exhaustion of L. polyactis stocks5,36, official statistics showed that the annual catch remains high. The L. polyactis production broke through 150,000 tons in 1995, and was above 300,000 tons after 2005. There is likely to have a large offshore stock of L. polyactis, which gradually joined the catch under increasing fishing efforts offshore. Furthermore, the L. polyactis stocks can be resilience to high pressure for several reasons: (1) its miscellaneous diet makes them be able to receive sufficient food sources; (2) size and age at sexual maturity reduced37,38; and (3) the over consumption of top predators relieves the prey pressure on middle trophic level species, such as L. polyactis, snappers, and flatfishes. A good job is the difficulties of artificial propagation and seedling breeding of small yellow croaker were broken for the first time in 201539 and the whole artificial cultivation was successfully realized in 2020 (https://www.chinanews.com.cn/cj/2020/07-02/9227715.shtml), which would effectively alleviate the market demand and wild stock sustain of small yellow croaker.Pelagic small fish stocks may not recovery quickly as early cognitionSmall pelagic fishes enjoy assembling in large schools of tens of thousands of individuals, and are more vulnerable to predators. Species S. sagax mainly filter plankton with a low trophic level about 2.8. It spawns in May–June, with high fecundity (an absolute fecundity of 30,000–100,000 pelagic eggs) and fast growth, and has short generation time of 1.4 years40. S. sagax shows strong phototropy, and can be caught using light purse seine, gill net, and fixed net fishing at night41,42.In 1989, the biomass of S. sagax was about twice of Bmsy. With the decreasing capture production of traditional economic fishes, S. sagax became a target species using specific fishing methods43, resulted in catch increase accompanied with B/Bmsy decline into a state of extremely unhealthy in 2019. Recovery of small pelagic species stocks would be delayed by the total catch control policy, mainly because the removal of large numbers of predator species left more opportunities for their feeding objects44. Resource rebuilding of S. sagax was not as quick as expected, as small pelagic species had to endure increasing predation pressure from the recovery of high-trophic species under the total catch control. At 1.0 Fmsy scenario, B/Bmsy would be only 0.88 by 2030, in need of a longer period to healthy state.Well-planned restocking can enhance resource recoverySwimming crab P. trituberculatus has high reproductive capacity, with a female can release two to three batches of eggs during a breeding season, and a batch contains about 1–6 million eggs45. Under the complementary of existing management measures and restocking programmes, the production of P. trituberculatus was kept in a certain amount close to a healthy state, and there is not an urgent need for its stock rebuilding. Since the 1990s, restocking of hatchery-produced larvae of P. trituberculatus has been promoted in coastal waters of China. Large-scaled restocking programmes were documented: 33 million larvae were released into the Yellow Sea by Shandong Province in June 2013 (http://hyj.shandong.gov.cn/xwzx/sjdt/201311/t20131120_507389.html); 50.3 million larvae with carapace width over 6 mm were released in the northern Yellow Sea by Liaoning Province in June 2020 (http://nync.ln.gov.cn/fwzx/zxdt/202007/t20200707_3902016.html); 16.1 million larvae were released into the East China Sea by Daishan County of Zhejiang Province in June 2021 (http://www.daishan.gov.cn/art/2021/6/8/art_1383064_59012675.html). What should be of concern is when, where, and how many seedlings are released46,47,48, to maximumly utilize the environmental resources without encroaching on the benefits of other species.Short-living species can be resilience to overfishingThe main cephalopod species in Chinese fisheries are Sepiella maindroni, mainly distributes in the East China Sea35 and Sepia esculenta, mainly distributes in the Bohai Sea, the Yellow Sea and the East China Sea49. As a 1-year lifespan species with fast growth rate, S. maindroni forms spawning migration from deep water to shallow nearshore bays in spring, partly within the fishing moratorium period. Due to the positive phototaxis, the cuttlefishes can be captured by light seining. Sepiella esculenta was the most important cephalopod economically in the northern coastal seas and one of the four major fisheries in the Bohai Sea and the Yellow Sea until the 1970s50. The abundance of this species has been greatly reduced with continuous fishing pressures and dwindling spawning grounds51.Total catch control and fishing moratorium showed significant output on the short lifespan cuttlefishes. Without the implementation of the “zero-growth” policy, the cuttlefishes resources would have been exhausted by 2015 and impossible to rebuild. According to the current state of resources, by 2030 the cuttlefish stocks can be recovered under the 1.0 Fmsy scenario. Moreover, the extent of cuttlefishes stock recovery relies on food supply.Ways to sustain fisheriesThe conflict between rising demand for fishery products and declining resources under multiple pressures including overfishing, climate change, and marine pollution has put heavy pressures at a global scale52. Chinese government has undertaken serious reforms to effectively replan the fishery industry.The effective recovery and rational utilization of resources depend on the support by sufficient reliable data. China started fishery statistics right after the foundation of the People’s Republic of China, completed by MOA (1949–2017) and MARA (since 2018). However, the statistical dataset has been questioned internationally53. According to the explanation by FAO54, before 2000s, especially from 1979 to the late 1990s, as the central government raced to meet the increasing demand for seafood and to grow the domestic production, the local governments had frequently overreported their local catch. In addition, fishermen may falsely claim to increase their production for surplus compensation, after the government introduced fishing subsidies. On the contrary, the production might have been underreported since the early 2000s55,56, which could be attributed to the existence of a large number of “black ships” (fishing vessels without relevant legal permits). Moreover, the lack of professionals in the early period and inaccurate knowledge of species identification by fishermen also lead to data uncertainty. Reasonable fisheries data should be consistent with the species functional traits and life history characteristics. However, in the actual fishing activities, the intentional and high-intensity selective fishing of species may greatly deviate the catch data from the data predicted by models. The Chinese government has been trying to improve the statistical system, including data coefficient adjustment, training of fishermen and professional, and supervision of statistical authorities5. In this study, selected objects are inshore species: the species are familiar to fishermen; the fishing vessel supervision is in place; the data collection is relatively rational and complete; all these are conducive to the reliability of the results.The zero-growth policy, which has been implemented since 1999, is an important measure in the history of marine fishery development and management in China. That is, the total catch of marine fisheries in the current year cannot be higher than that of the previous year. However, the “12th Five-year Plan” for national fishery development (2011–2015) issued by the Ministry of Agriculture canceled the mandatory targets of controlling the production but to encourage more catches of marine fisheries (http://www.moa.gov.cn/gk/ghjh_1/201110/t20111017_2357716.htm). In 2013, the State Council published the first state-level marine fishery development document as “Several Advices on Promoting Marine Sustainable and Healthy Development”, incorporating marine fishery development into the strategy of building a maritime power (http://www.gov.cn/zwgk/2013-06/25/content_2433577.htm). This policy shift was clearly reflected in the significant increase in the national annual catch from 12 to 14 MT. Until the “13th Five-year Plan” for national fishery development (2016–2020) issued in 2016, the zero-even negative-growth policy was revalidated, and the volume of annual output control was clearly proposed as 8–10 MT57, which was determined by multiplying the fishing coefficient by the total stock size derived from the assessment of surveys on the zoning of fisheries and the supplementary survey of marine biological resources in the exclusive economic zone and the continental shelf7. To achieve the target of keeping fishing capacity at a high level of sustainability, significant reductions in fishing pressures over a period of time are required, as well as rational updates of control policies.Many policies were introduced together or around the same time as the “zero-growth” policy, such as summer fishing moratorium, fishing license system, and fishing fuel subsides. However, the achievements are far from satisfactory. The fishing fuel subsidy policy together with the license system induced the direct fishing vessel construction boom which resulted in fewer but bigger and more powerful fishing vessels. Fishing moratorium is the most promising policy, by leaving enough time and space for fish to successfully reproduce. However, the truth is that, right after the fishing closure season, almost all fishing vessels immediately rush into the sea and fishermen try their best to fish as much as possible within the gears and engine power permission of their fishing licenses, attempting to earn a year’s income in a short period of 2–4 months. As a result of such high fishing effort, the achievements of seasonal fishing bans were largely offset and resource densities fell to low levels after autumn. The number of legally binding standards for mesh size is not enough, only 6 at present of at least 40 fishing target species and over 10 fishing gears, leaving many fishing gears and fish species outside the regulation of existing standards6,58. Ideally, standards of mesh size should be updated corresponding to the changes of species traits, however, it is a challenge because the main fishing mode is multiple species fishery by bottom trawling. Moreover, species in China seas are diverse, and the spawning period of different species may not fall into the fishing closure season5. The lack of specificity to sufficiently cover all the species may result an unbalance of community composition. Another system “Double Control” aims to limit both the numbers of fishing vessels and the total power. Unfortunately, the inspections of fishing vessels and their power are not very strict, due to the need of developing local economy and guaranteeing the fishermen’s income, e.g., under a nominal power mask the low-power engines have been replaced by high-power engines, some fishing vessels do not have the fishing licenses28. The limitation of the license number and engine power also stimulate the technological improvement for more catch7.The structure adjustment of fisheries composition is the main management measure at present. The high degree of self-sufficiency in fishery products in China has been achieved through overfishing of domestic fishery resources, resulting in the rapid depletion of fisheries in China’s coastal waters59. Aquaculture, accounting for more than 70% of China’s total fisheries production2, is identified as a successful way. Accompanying by aquaculture development, a series of problems also arise, particularly, the demand of low-value/trash fish and fish meal that significantly drives further expansion of capture fisheries60. Cooperation with other countries to promote regional aquaculture may be an alternative way to meeting global growing demand for seafood and combating overfishing61,62. Seeking resources from the high seas and EEZs of other countries is also a choice, of course, on the premise of taking full account of ecology, maritime, and food security of other countries63,64,65.In addition, this study pointed out a new focus for fisheries management, in which differences in species biological traits, including species vulnerability, population multiplication, and resilience to environmental pressures, should be given full consideration. On this basis, more detailed and targeted management schemes are supposed to propose to achieve the dual purpose of recoverable fisheries resource and balanced species composition, so as to become a truly sustainable fishery. In short, the effective implementation of various management measures is an indispensable guarantee. More

Lenton, T. & Watson, A. J. Revolutions That Made the Earth. (Oxford University Press, 2011).Lovelock, J. The Ages of Gaia: A Biography of Our Living Earth. (Oxford University Press, USA, 2000).Falkowski, P. G. The rise of oxygen over the past 205 million years and the evolution of large placental mammals. Science 309, 2202–2204 (2005).Article 
PubMed 

Google Scholar 
Holland, H. D. The oxygenation of the atmosphere and oceans. Philos. Trans. R. Soc. B: Biol. Sci. 361, 903–915 (2006).Article 

Google Scholar 
Lenton, T. M. Fire feedbacks on atmospheric oxygen. In Fire phenomena and the Earth system: an interdisciplinary guide to fire science (ed. Belcher, C. M.) 289–308 (John Wiley & Sons, 2013).Belcher, C. M., Yearsley, J. M., Hadden, R. M., McElwain, J. C. & Rein, G. Baseline intrinsic flammability of Earth’s ecosystems estimated from paleoatmospheric oxygen over the past 350 million years. Proc. Natl Acad. Sci. 107, 22448–22453 (2010).Article 
PubMed 
PubMed Central 

Google Scholar 
Cope, M. J. & Chaloner, W. G. Fossil charcoal as evidence of past atmospheric composition. Nature 283, 647–649 (1980).Article 

Google Scholar 
Watson, A. J. Consequences for the biosphere of forest and grassland fires. (University of Reading, 1978).Belcher, C. M. & McElwain, J. C. Limits for combustion in low O2 redefine paleo atmospheric predictions for the Mesozoic. Science 321, 1197–1200 (2008).Article 
PubMed 

Google Scholar 
Wildman, R. A., Hickey, L. J., Dickinson, M. B. & Wildman, C. B. Burning of forest materials under late Paleozoic high atmospheric oxygen levels. Geology 32, 457–460 (2004).Kump, L. R. The rise of atmospheric oxygen. Nature 451, 277–278 (2008).Article 
PubMed 

Google Scholar 
Glasspool, I. J., Edwards, D. & Axe, L. Charcoal in the Silurian as evidence for the earliest wildfire. Geology 32, 381–383 (2004).Article 

Google Scholar 
Bowman, D. M. et al. Fire in the Earth system. Science 324, 481–484 (2009).Article 
PubMed 

Google Scholar 
Quintiere, J. G. Principles of Fire Behaviour. (CRC Press Boca Raton, 1998).Pyne, S. J., Andrews, P. L. & Laven, R. D. Introduction to wildland fire. (John Eiley & Sons, Inc., 1996).Jones, T. P. & Chaloner, W. G. Fossil charcoal, its recognition and palaeoatmospheric significance. Palaeogeogr. Palaeoclimatol. Palaeoecol. 97, 39–50 (1991).Article 

Google Scholar 
Glasspool, I. J. & Scott, A. C. Phanerozoic concentrations of atmospheric oxygen reconstructed from sedimentary charcoal. Nat. Geosci. 3, 627–630 (2010).Article 

Google Scholar 
Belcher, C. M., Collinson, M. E. & Scott, A. C. Fire phenomena and the Earth system: an interdisciplinary guide to fire science. In A 450‐Million‐Year History of Fire 229–249 (Wiley Online Library, 2013).Berner, R. A. & Landis, G. P. Chemical analysis of gaseous bubble inclusions in amber; the composition of ancient air? Am. J. Sci. 287, 757–762 (1987).Article 

Google Scholar 
Lane, N. Oxygen: The Molecule that Made the World. (Oxford University Press, 2002).Hopfenberg, H. B. et al. Is the air in amber ancient? Science 241, 717–721 (1988).Article 
PubMed 

Google Scholar 
Carpenter, F. M. Studies on Carboniferous insects from Commentry, France; Part I. Introduction and families Protagriidae, Meganeuridae, and Campylopteridae. Bull. Geol. Soc. Am. 54, 527–554 (1943).Article 

Google Scholar 
Carpenter, F. M. Studies on Carboniferous insects from Commentry, France: Part II. The Megasecoptera. J. Paleontol. 25, 336–355 (1951).
Google Scholar 
Whyte, M. A. A gigantic fossil arthropod trackway. Nature 438, 576–576 (2005).Article 
PubMed 

Google Scholar 
Carroll, R. L. Vertebrate Paleontology and Evolution. (Freeman, 1988).Graham, J. B., Aguilar, N. M., Dudley, R. & Gans, C. Implications of the late Palaeozoic oxygen pulse for physiology and evolution. Nature 375, 117–120 (1995).Article 

Google Scholar 
Harrison, J. F., Kaiser, A. & VandenBrooks, J. M. Atmospheric oxygen level and the evolution of insect body size. Proc. R. Soc. B: Biol. Sci. 277, 1937–1946 (2010).Article 

Google Scholar 
Hetz, S. K. & Bradley, T. J. Insects breathe discontinuously to avoid oxygen toxicity. Nature 433, 516–519 (2005).Article 
PubMed 

Google Scholar 
Watson, A., Lovelock, J. E. & Margulis, L. Methanogenesis, fires and the regulation of atmospheric oxygen. Biosystems 10, 293–298 (1978).Article 
PubMed 

Google Scholar 
Watson, A. J. & Lovelock, J. E. The dependence of flame spread and probability of ignition on atmospheric oxygen: an experimental investigation. In Fire phenomena and the Earth system: an interdisciplinary guide to fire science 273–287 (John Wiley & Sons, 2013).Thonicke, K., Venevsky, S., Sitch, S. & Cramer, W. The role of fire disturbance for global vegetation dynamics: coupling fire into a Dynamic Global Vegetation Model. Glob. Ecol. Biogeogr. 10, 661–677 (2001).Article 

Google Scholar 
Benson, R. P., Roads, J. O. & Weise, D. R. Climatic and weather factors affecting fire occurrence and behavior. Dev. Environ. Sci. 8, 37–59 (2008).
Google Scholar 
Babrauskas, V. Effective heat of combustion for flaming combustion of conifers. Can. J. For. Res. 36, 659–663 (2006).Article 

Google Scholar 
Madrigal, J., Guijarro, M., Hernando, C., Diez, C. & Marino, E. Effective heat of combustion for flaming combustion of Mediterranean forest fuels. Fire Technol. 47, 461–474 (2011).Article 

Google Scholar 
Rivera, J., de, D., Davies, G. M. & Jahn, W. Flammability and the heat of combustion of natural fuels: a review. Combust. Sci. Technol. 184, 224–242 (2012).Article 

Google Scholar 
Dibble, A. C., White, R. H. & Lebow, P. K. Combustion characteristics of north-eastern USA vegetation tested in the cone calorimeter: invasive versus non-invasive plants. Int. J. Wildland Fire 16, 426–443 (2007).Article 

Google Scholar 
Stein, W. E. et al. Mid-Devonian Archaeopteris roots signal revolutionary change in earliest fossil forests. Curr. Biol. 30, 421–431.e2 (2020).Article 
PubMed 

Google Scholar 
Lenton, T. M. & Watson, A. J. Redfield revisited: 2. What regulates the oxygen content of the atmosphere? Glob. Biogeochem. Cycles 14, 249–268 (2000).Article 

Google Scholar 
Berner, R. A. The Phanerozoic Carbon Cycle: CO2 and O2. (Oxford University Press on Demand, 2004).Berner, R. A. GEOCARBSULF: a combined model for Phanerozoic atmospheric O2 and CO2. Geochimica et. Cosmochimica Acta 70, 5653–5664 (2006).Article 

Google Scholar 
Berner, R. A. GEOCARB II: A revised model of atmospheric CO2 over phanerozoic time. Am. J. Sci. 294, 56–91 (1994).Bergman, N. M., Lenton, T. M. & Watson, A. J. COPSE: a new model of biogeochemical cycling over Phanerozoic time. Am. J. Sci. 304, 397–437 (2004).Article 

Google Scholar 
Lenton, T. M., Daines, S. J. & Mills, B. J. COPSE reloaded: an improved model of biogeochemical cycling over Phanerozoic time. Earth-Sci. Rev. 178, 1–28 (2018).Article 

Google Scholar 
Mills, B. J., Donnadieu, Y. & Goddéris, Y. Spatial continuous integration of Phanerozoic global biogeochemistry and climate. Gondwana Res. 100, 73–86 (2021).Article 

Google Scholar 
Kump, L. R. Terrestrial feedback in atmospheric oxygen regulation by fire and phosphorus. Nature 335, 152–154 (1988).Article 

Google Scholar 
Holland, H. D. The Chemical Evolution of the Atmosphere and Oceans. vol. 2 (Princeton University Press, 2020).Lasaga, A. C. & Ohmoto, H. The oxygen geochemical cycle: dynamics and stability. Geochimica et. Cosmochimica Acta 66, 361–381 (2002).Article 

Google Scholar 
Van Cappellen, P. & Ingall, E. D. Redox stabilization of the atmosphere and oceans by phosphorus-limited marine productivity. Science 271, 493–496 (1996).Article 
PubMed 

Google Scholar 
Belcher, C. M. et al. The rise of angiosperms strengthened fire feedbacks and improved the regulation of atmospheric oxygen. Nat. Commun. 12, 503 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Belcher, C. M., Yearsley, J. M., Hadden, R. M., McElwain, J. C. & Rein, G. Baseline intrinsic flammability of Earth’s ecosystems estimated from paleoatmospheric oxygen over the past 350 million years. Proc. Natl Acad. Sci. 107, 22448–22453 (2010).Article 
PubMed 
PubMed Central 

Google Scholar 
Berner, R. A. & Canfield, D. E. A new model for atmospheric oxygen over Phanerozoic time. Am. J. Sci. 289, 333–361 (1989).Article 
PubMed 

Google Scholar 
Lenton, T. M. The role of land plants, phosphorus weathering and fire in the rise and regulation of atmospheric oxygen. Glob. Change Biol. 7, 613–629 (2001).Article 

Google Scholar 
Royer, D. L., Donnadieu, Y., Park, J., Kowalczyk, J. & Godderis, Y. Error analysis of CO2 and O2 estimates from the long-term geochemical model GEOCARBSULF. Am. J. Sci. 314, 1259–1283 (2014).Article 

Google Scholar 
Berner, R. A. Inclusion of the weathering of volcanic rocks in the GEOCARBSULF model. Am. J. Sci. 306, 295–302 (2006).Article 

Google Scholar 
Keeley, J. E., Pausas, J. G., Rundel, P. W., Bond, W. J. & Bradstock, R. A. Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci. 16, 406–411 (2011).Article 
PubMed 

Google Scholar 
Pausas, J. G. & Keeley, J. E. A burning story: the role of fire in the history of life. BioScience 59, 593–601 (2009).Article 

Google Scholar 
Bond, W. J., Woodward, F. I. & Midgley, G. F. The global distribution of ecosystems in a world without fire. N. Phytol.t 165, 525–538 (2005).Article 

Google Scholar 
Forkel, M. et al. Emergent relationships with respect to burned area in global satellite observations and fire-enabled vegetation models. Biogeosciences 16, 57–76 (2019).Article 

Google Scholar 
Lucht, W., Schaphoff, S., Erbrecht, T., Heyder, U. & Cramer, W. Terrestrial vegetation redistribution and carbon balance under climate change. Carbon Balance Manag. 1, 1–7 (2006).Article 

Google Scholar 
Wu, C. et al. Historical and future global burned area with changing climate and human demography. One Earth 4, 517–530 (2021).Article 

Google Scholar 
Thonicke, K. et al. The influence of vegetation, fire spread and fire behaviour on biomass burning and trace gas emissions: results from a process-based model. Biogeosciences 7, 1991–2011 (2010).Article 

Google Scholar 
Sitch, S. et al. Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model. Glob. Change Biol. 9, 161–185 (2003).Article 

Google Scholar 
Lovelock, J. E. Gaia: A New Look at Life on Earth. (Oxford Paperbacks, 2000).Lasslop, G. et al. Global ecosystems and fire: Multi‐model assessment of fire‐induced tree‐cover and carbon storage reduction. Glob. Change Biol. 26, 5027–5041 (2020).Article 

Google Scholar 
Quan, X. et al. Global fuel moisture content mapping from MODIS. Int. J. Appl. Earth Obs. Geoinf. 101, 102354 (2021).
Google Scholar 
Collinson, M. E. et al. Palynological evidence of vegetation dynamics in response to palaeoenvironmental change across the onset of the Paleocene‐Eocene Thermal Maximum at Cobham, Southern England. Grana 48, 38–66 (2009).Article 

Google Scholar 
Feurdean, A. & Vasiliev, I. The contribution of fire to the late Miocene spread of grasslands in eastern Eurasia (Black Sea region). Sci. Rep. 9, 1–7 (2019).Article 

Google Scholar 
Hollaar, T. P. et al. Wildfire activity enhanced during phases of maximum orbital eccentricity and precessional forcing in the Early Jurassic. Commun. Earth Environ. 2, 1–12 (2021).Article 

Google Scholar 
Zelitch, I. Photosynthesis, Photorespiration, and Plant Productivity. (Elsevier, 2012).Björkman, O. The effect of oxygen concentration on photosynthesis in higher plants. Physiol. Plant. 19, 618–633 (1966).Article 

Google Scholar 
Berner, R. A. & Kothavala, Z. GEOCARB III: a revised model of atmospheric CO2 over Phanerozoic time. Am. J. Sci. 301, 182–204 (2001).Article 

Google Scholar 
Baker, S. J., Hesselbo, S. P., Lenton, T. M., Duarte, L. V. & Belcher, C. M. Charcoal evidence that rising atmospheric oxygen terminated Early Jurassic ocean anoxia. Nat. Commun. 8, 1–7 (2017).Article 

Google Scholar 
Pfeiffer, M., Spessa, A. & Kaplan, J. O. A model for global biomass burning in preindustrial time: LPJ-LMfire (v1.0). Geosci. Model Dev. 6, 643–685 (2013).Article 

Google Scholar 
Cohen, J. D. The national fire-danger rating system: basic equations. vol. 82 (US Department of Agriculture, Forest Service, Pacific Southwest Forest and …, 1985). More

McNeely JA. editor. Global strategy on invasive alien species. IUCN (2001).Perrings, C. et al. Biological invasion risks and the public good: An economic perspective. Conserv. Ecol. 6, 1 (2002).
Google Scholar 
Taylor, B. W. & Irwin, R. E. Linking economic activities to the distribution of exotic plants. Proc. Natl. Acad. Sci. U.S.A. 101, 17725–17730 (2004).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Moore, B. A. Alien invasive species: Impacts on forests and forestry—A review (Forestry Department and Forest Resource Division FAO, FAO Corporate Document Repository, 2005).
Google Scholar 
McBeath, J. H. & McBeath, J. Invasive Species and Food Security 157–176 (In Environmental Change and Food Security in China. Springer, 2010).
Google Scholar 
Ziska, L. H., Blumenthal, D. M., Runion, G. B., Hunt, E. R. & Diaz-Soltero, H. Invasive species and climate change: An agronomic perspective. Clim. Change. 105, 13–42 (2011).Article 
ADS 

Google Scholar 
Doherty, T. S., Glen, A. S., Nimmo, D. G., Ritchie, E. G. & Dickman, C. R. Invasive predators and global biodiversity loss. Proc. Natl. Acad. Sci. U.S.A. 113, 11261–11265 (2016).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Otero, R. P., Vázquez, J. P. M. & Del Estal, P. Detección de la psila africana de los cítricos, Trioza erytreae (Del Guercio, 1918) (Hemiptera: Psylloidea: Triozidae), en la Península Ibérica. Arquivos Entomolóxicos 13, 119–122 (2015).
Google Scholar 
van den Berg, M. A., Deacon, & V. E.,. Dispersal of the citrus psylla, Trioza erytreae (Hemiptera: Triozidae), in the absence of its host plants. Phytophylactica 20, 361–368 (1988).
Google Scholar 
CABI. Trioza erytreae. In: Invasive Species Compendium. Wallingford, UK: CAB International. www.cabi.org/isc. (2021).Lounsbury, C. P. Psyllidae or jumping plant lice in Report of the Government Entomologist for the year 1896. Cape of Good Hope, South Africa, (Unpublished report), 115–118 (1897).Ruíz-Rivero, O. et al. Insights into the origin of the invasive populations of Trioza erytreae in Europe using microsatellite markers and mtDNA barcoding approaches. Sci. Rep. 11, 1–15 (2021).Article 

Google Scholar 
Benhadi-Marín, J., Fereres, A. & Pereira, J.A. Potential areas of spread of Trioza erytreae over mainland Portugal and Spain. J. Pest Sci.1–12 (2021).Bové, J.M. Huanglongbing: a destructive, newly-emerging, century-old disease of citrus. Plant Pathol. 7–37 (2006).Laštuvka, Z. Climate change and its possible influence on the occurrence and importance of insect pests. Plant Prot. Sci. 45, S53–S62 (2009).Article 

Google Scholar 
Thomson, L. J., Macfadyen, S. & Hoffmann, A. A. Predicting the effects of climate change on natural enemies of agricultural pests. Biol. Control. 52, 296–306 (2010).Article 

Google Scholar 
Bajwa, A.A., Farooq, M., Al-Sadi, A.M., Nawaz, A., Jabran, K. & Siddique, K.H. Impact of climate change on biology and management of wheat pests. J. Crop Prot. 105304 (2020).Hamann, E., Blevins, C., Franks, S. J., Jameel, M. I. & Anderson, J. T. Climate change alters plant–herbivore interactions. New Phytol. 229, 1894–1910 (2021).Article 
CAS 
PubMed 

Google Scholar 
Cornelissen, T. Climate change and its effects on terrestrial insects and herbivory patterns. Neotrop. Entomol. 40, 155–163 (2011).Article 
CAS 
PubMed 

Google Scholar 
Raffa, K. F. et al. Responses of tree-killing bark beetles to a changing climate. Clim. Change Insect Pests. 7, 173–201 (2015).Article 

Google Scholar 
Cocuzza, G. E. M. et al. A review on Trioza erytreae (African citrus psyllid), now in mainland Europe, and its potential risk as vector of huanglongbing (HLB) in citrus. J. Pest Sci. 90, 1–17 (2017).Article 

Google Scholar 
Vector of citrus greening disease. Aidoo, O. F., Tanga, C. M., Azrag, A. G., Mohamed, S. A., Khamis, F. M., Rasowo, B. A. … & Borgemeister, C. Temperature-based phenology model of African citrus triozid (Trioza erytreae Del Guercio). J. Appl. Entomol. 146, 1–2 (2021).
Google Scholar 
Catling, H. D., The bionomics of the South African citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: PsyUidae), 1. The influence of the flushing rhythm of citrus and factors which regulate flushing. J. Entomol. Soc. S. Afr. 32, 191–208 (1969).Green, G. C. E., & Catling, H. D. “Weather-induced mortality of the citrus psylla, Trioza erytreae (Del Guercio)(Homoptera: Psyllidae), a vector of greening virus, in some citrus producing areas of southern Africa.” Agric. Meteorol. 8, 305–317(1971).Vicente-Serrano, S. M., González-Hidalgo, J. C., de Luis, M. & Raventós, J. Drought patterns in the Mediterranean area: The Valencia region (eastern Spain). Clim. Res. 26, 5–15 (2004).Article 

Google Scholar 
Millán, M. M., Estrela, M. J. & Miró, J. Rainfall components: variability and spatial distribution in a Mediterranean Area (Valencia Region). J. Clim. 18, 2682–2705 (2005).Article 
ADS 

Google Scholar 
Srivastava, V., Lafond, V. & Griess, V.C. Species distribution models (SDM): applications, benefits and challenges in invasive species management. CAB Rev. 14(10.1079) (2019).Halsch, C.A., Shapiro, A.M., Fordyce, J.A., Nice, C.C., Thorne, J.H., Waetjen, D.P. & Forister, M.L. Insects and recent climate change. Proc. Natl. Acad. Sci. U.S.A. 118 (2021).Elith, J. & Leathwick, J. R. Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697 (2009).Article 

Google Scholar 
Lobo, J. M., Jiménez-Valverde, A. & Hortal, J. The uncertain nature of absences and their importance in species distribution modelling. ECOGEG 33, 103–114 (2010).
Google Scholar 
Guisan, A., Thuiller, W. & Zimmermann, N.E. Habitat suitability and distribution models: with applications in R. Cambridge University Press. (2017).de la Vega, G. J. & Corley, J. C. Drosophila suzukii (Diptera: Drosophilidae) distribution modelling improves our understanding of pest range limits. Int. J. Pest Manag. 65, 217–227 (2019).Article 

Google Scholar 
Tavanpour, T., Sarafrazi, A., Mehrnejad, M.R. & Imani, S. Distribution modelling of Acrosternum spp. (Hemiptera: Pentatomidae) in south of Iran. Biologia, 74, 1627–1635 (2019).Barton, M. G. & Terblanche, J. S. Predicting performance and survival across topographically heterogeneous landscapes: the global pest insect Helicoverpa armigera (H übner, 1808) (L epidoptera: N octuidae). Austral. Entomol. 53, 249–258 (2014).Article 

Google Scholar 
Kearney, M. & Porter, W. P. Mechanistic niche modelling: Combining physiological and spatial data to predict species’ ranges. Ecol. Lett. 12, 334–350 (2009).Article 
PubMed 

Google Scholar 
Thomas, C. D. et al. Extinction risk from climate change. Nature 427, 145–148 (2004).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Shabani, F., Kumar, L. & Ahmadi, M. A comparison of absolute performance of different correlative and mechanistic species distribution models in an independent area. Ecol. Evol. 6, 5973–5986 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Kearney, M. R., Wintle, B. A. & Porter, W. P. Correlative and mechanistic models of species distribution provide congruent forecasts under climate change. Conserv. Lett. 3, 203–213 (2010).Article 

Google Scholar 
Moran, V. C. & Blowers, J. R. On the biology of the South African citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: Psyllidae). J. Entomol. Soc. S. Afr. 30, 96–106 (1967).
Google Scholar 
Samways, M.J. & Manicom, B.Q. Immigration, frequency distributions and dispersion patterns of the psyllid Trioza erytreae (Del Guercio) in a citrus orchard. J. Appl. Ecol. 463–472 (1983).Pérez-Rodríguez, J. et al. Classical biological control of the African citrus psyllid Trioza erytreae, a major threat to the European citrus industry. Sci. Rep. 9, 1–11 (2019).Article 

Google Scholar 
Aidoo, O. F. et al. Host suitability and feeding preference of the African citrus triozid Trioza erytreae Del Guercio (Hemiptera: Triozidae), natural vector of “Candidatus Liberibacter africanus”. J. Appl. Entomol. 143, 262–270 (2019).Article 

Google Scholar 
Moran, V. C. Preliminary observations on the choice of host plants by adults of the citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: Psyllidae). J. Entomol. Soc. S. Afr. 31, 403–410 (1968).
Google Scholar 
van den Berg, M. A., Deacon, V. E. & Thomas, C.D. Ecology of the citrus psylla, Trioza erytreae (Hemiptera: Triozidae). 3. Mating, fertility and oviposition. Phytophylactica. 23, 195–200 (1991).Khamis, F. M. et al. DNA barcode reference library for the African citrus triozid, Trioza erytreae (Hemiptera: Triozidae): Vector of African citrus greening. J. Econ. Entomol. 110, 2637–2646 (2017).Article 
CAS 
PubMed 

Google Scholar 
Aidoo, O. F. et al. The African citrus triozid Trioza erytreae Del Guercio (Hemiptera: Triozidae): temporal dynamics and susceptibility to entomopathogenic fungi in East Africa. Int. J. Trop. Insect Sci. 41, 563–573 (2021).Article 

Google Scholar 
Rasowo, B. A. et al. Diversity and phylogenetic analysis of endosymbionts from Trioza erytreae (Del Guercio) and its parasitoids in Kenya. J. Appl. Entomol. 145, 104–116 (2021).Article 

Google Scholar 
Espinosa-Zaragoza, S., Aguirre-Medina, J. F. & López-Martínez, V. Does the African Citrus psyllid, Trioza erytreae (Del Guercio) (Hemiptera: Triozidae), Represent a phytosanitary threat to the citrus industry in Mexico?. Insects. 12, 450 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Aidoo, O.F., Tanga, C.M., Mohamed, S.A., Khamis, F.M., Baleba, S.B., Rasowo, B.A., Ambajo, J., Sétamou, M., Ekesi, S. & Borgemeister, C. Detection and monitoring of ‘Candidatus’ Liberibacter spp. vectors: African citrus triozid Trioza erytreae Del Guercio (Hemiptera: Triozidae) and Asian citrus psyllid Diaphorina citri Kuwayama (Hemiptera: Liviidae) in citrus groves in East Africa. Agric. For. Entomol. 22, 401–409 (2020a).Urbaneja-Bernat, P., Hernández-Suárez, E., Tena, A. & Urbaneja, A. Preventive measures to limit the spread of Trioza erytreae (Del Guercio) (Hemiptera: Triozidae) in mainland Europe. J. Appl. Entomol. 144, 553–559 (2020).Article 

Google Scholar 
Aidoo, O. F. et al. Size and shape analysis of Trioza erytreae Del Guercio (Hemiptera: Triozidae), vector of citrus huanglongbing disease. Pest Manag. Sci. 75, 760–771 (2019).Article 
CAS 
PubMed 

Google Scholar 
Arenas-Arenas, F. J., Duran-Vila, N., Quinto, J. & Hervalejo, A. Geographic spread and inter-annual evolution of populations of Trioza erytreae in the Iberian Peninsula. Plant Pathol. 101, 1151–1157 (2019).Article 

Google Scholar 
Kalyebi, A. et al. Detection and identification of etiological agents (Liberibacter spp.) associated with citrus greening disease in Uganda. J. Agric. Sci. 16, 43–54 (2015).
Google Scholar 
Kyalo Richard., Abdel-Rahman, E.M., Mohamed, S.A., Ekesi, S., Borgemeister, C. & Landmann, T. Importance of remotely-sensed vegetation variables for predicting the spatial distribution of African citrus triozid (Trioza erytreae) in Kenya. ISPRS Int. J. Geoinf. 7, 429 (2018).Benhadi-Marín, J., Fereres, A. & Pereira, J. A. A model to predict the expansion of Trioza erytreae throughout the Iberian Peninsula using a pest risk analysis approach. Insects. 11, 576 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Moran, V. C. The development of the citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: Psyllidae), on Citrus limon and four indigenous host plants. J. Entomol. Soc. S. Afr. 31, 391–402 (1986).
Google Scholar 
Tamesse, J. L. Key for identification of the Hymenopteran parasitoids of the African citrus psylla Trioza erytreae Del Guercio (Hemiptera: Triozidae) in Cameroon. Afr. J. Agric. Res. 4, 085–091 (2009).
Google Scholar 
Hailu, T. & Wakgari, M. Distribution and damage of African citrus psyllids (Trioza erytreae) in Casimiroa edulis producing areas of the eastern zone of Ethiopia. Int. J. Environ. Agric. Biotech. 4, 741–750 (2019).
Google Scholar 
Urbaneja-Bernat, P. et al. Host range testing of Tamarixia dryi (Hymenoptera: Eulophidae) sourced from South Africa for classical biological control of Trioza erytreae (Hemiptera: Psyllidae) in Europe. Biol. Control. 135, 110–116 (2019).Article 

Google Scholar 
Hernández-Suárez, E., Pérez-Rodríguez, J., Suárez-Méndez, L., Urbaneja-Bernat, P., Rizza, R., Siverio, F., Piedra-Buena, A., Urbaneja, A. &Tena, A.. Control de Trioza erytreae en las Islas Canarias por el parasitoide Tamarixia dryi. Phytoma España. La revista profesional de sanidad vegetal. 28–32 (2021).Molina, P., Martínez-Ferrer, M. T., Campos-Rivela, J. M., Riudavets, J. & Agustí, N. Development of a PCR-based method for the screening of potential predators of the African citrus psyllid Trioza erytreae (Del Guercio). Biol. Control. 160, 104661 (2021).Article 
CAS 

Google Scholar 
Kumar, S., Neven, L. G., & Yee, W. L. Evaluating correlative and mechanistic niche models for assessing the risk of pest establishment. Ecosphere. 5, (2014).Kriticos, D. J. et al. The potential distribution of invading Helicoverpa armigera in North America: Is it just a matter of time?. PLoS ONE 10, e0119618 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Sutherst, R. W., Maywald, G. F. & Bourne, A. S. Including species interactions in risk assessments for global change. Glob. Chang. Biol. 13, 1843–1859 (2007).Article 
ADS 

Google Scholar 
Shabani, F., Kumar, L. & Esmaeili, A. Use of CLIMEX, land use and topography to refine areas suitable for date palm cultivation in Spain under climate change scenarios. J. Earth Sci. Clim. Change. 4, 145 (2013).
Google Scholar 
Silva, R. S., Kumar, L., Shabani, F. & Picanço, M. C. Assessing the impact of global warming on worldwide open field tomato cultivation through CSIRO-Mk3•0 global climate model. J. Agric. Sci. 155, 407–420 (2016).Article 

Google Scholar 
Kriticos, D. J. et al. CliMond: Global high-resolution historical and future scenario climate surfaces for bioclimatic modelling. Methods Ecol. Evol. 3, 53–64 (2012).Article 

Google Scholar 
Catling, H. D. The bionomics of the South African citrus psylla, Trioza erytreae (Del Guercio) (Homoptera: Psyllidae) 3. The influence of extremes of weather on survival. J. Ecol. Soc. S. Afr. 32, 273–290 (1969).Aubert, B. Trioza erytreae Del Guercio and Diaphorina citri Kuwayama (Homoptera: Psylloidea), the two vectors of citrus greening disease: biological aspects and possible control strategy. Fruits 42, 149–162 (1987).
Google Scholar 
Gordon, H. B., Rotstayn, L. D., Mcgregor, J. L., Dix, M. R., Kowalczyk, E. A., O’farrell, S. P., Waterman, L. J., Hirst, A. C., Wilson, S. G., Collier, M. A., Watterson, I. G. & Elliott, T. I. The CSIRO Mk3 Climate System Model. CSIRO Atmospheric Research Technical Paper No. 60. Canberra: CSIRO. (2002).Van Vuuren, D. P. & Carter, T. R. Climate and socio-economic scenarios for climate change research and assessment: reconciling the new with the old. Clim. Change. 122, 415–429 (2013).Article 
ADS 

Google Scholar 
Fecher, B., Friesike, S. & Hebing, M. What drives academic data sharing?. PLoS ONE 10, e0118053 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Imker, H. J., Luong, H., Mischo, W. H., Schlembach, M. C. & Wiley, C. An examination of data reuse practices within highly cited articles of faculty at a research university. J. Acad. Librariansh. 47, 102369 (2021).Article 

Google Scholar 
Aidoo, O. F. et al. Distribution, degree of damage and risk of spread of Trioza erytreae (Hemiptera: Triozidae) in Kenya. J. Appl. Entomol. 143, 822–833 (2019).Article 

Google Scholar 
Mack, R.N., Simberloff, D., Mark Lonsdale, W., Evans, H., Clout, M. & Bazzaz, F.A. Biotic invasions: causes, epidemiology, global consequences, and control. Ecol. Appl. 10, 689–710 (2000).EPPO. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. https://gd.eppo.int/ (2021).Beattie, G.A.C., Holford, P., Mabberley, D.J., Haigh, A.M. and Broadbent, P. Australia and huanglongbing. Food & Fertilizer Technology Center. (2008).Beattie, G.A.C. & Barkley, P. Huanglongbing and its Vectors. A Pest Specific Contingency Plan for the Citrus and Nursery and Garden Industries (Version 2), February 2009. Horticulture Australia Ltd., Sydney (2009).Plant Biosecurity. Final pest risk analysis report for ‘Candidatus Liberibacter species’ and their vectors associated with Rutaceae. Department of Agriculture, Fisheries and Forestry, Canberra. (2011).Silva, R. S., Kumar, L., Shabani, F. & Picanço, M. C. Potential risk levels of invasive Neoleucinodes elegantalis (small tomato borer) in areas optimal for open-field Solanum lycopersicum (tomato) cultivation in the present and under predicted climate change. Pest. Manag. Sci. 73, 616–627 (2017).Article 
PubMed 

Google Scholar 
Santana, P. A., Kumar, L., Da Silva, R. S. & Picanço, M. C. Global geographic distribution of Tuta absoluta as affected by climate change. J. Pest Sci. 92, 1373–1385 (2019).Article 

Google Scholar 
da Graça, J. V. Citrus greening disease. Annu. Rev. Phytopathol. 29, 109–136 (1991).Article 

Google Scholar 
Li, W., Levy, L. & Hartung, J. S. Quantitative distribution of ‘Candidatus Liberibacter asiaticus’ in citrus plants with citrus huanglongbing. Phytopathology 99, 139–144 (2009).Article 
ADS 
PubMed 

Google Scholar 
Tatineni, S. et al. In Planta Distribution of ‘Candidatus Liberbacter asiaticus’ as revealed by Polymerase Chain Reaction (PCR) and Real-time PCR. Phytopathology 98, 592–599 (2008).Article 
CAS 
PubMed 

Google Scholar 
Aubert, B. Historical perspectives of HLB in Asia. In: International Research Conference on Huanglongbing; Proceedings of the Meeting (eds. Gottwald RT, Graham HJ) Orlando, Florida. 16–24 (2008).microscopy and microarray analysis. Kim, J, S., Sagaram, U.S., Burns, J.K., Li, J.L. & Wang, N. Response of sweet orange (Citrus sinensis) to ‘Candidatus Liberibacter asiaticus’ infection. Phytopathology 99, 50–57 (2009).Article 

Google Scholar 
EPPO. Trioza erytreae. EPPO datasheets on pests recommended for regulation (2022). Available online. https://gd.eppo.int.Ajene, I. J. et al. Habitat suitability and distribution potential of Liberibacter species (“Candidatus Liberibacter asiaticus” and “Candidatus Liberibacter africanus”) associated with citrus greening disease. Divers. Distrib. 26, 575–588 (2020).Article 

Google Scholar 
Manjunath, K. Á., Halbert, S. E., Ramadugu, C. H., Webb, S. U. & Lee, R. F. Detection of ‘Candidatus Liberibacter asiaticus’ in Diaphorina citri and its importance in the management of citrus huanglongbing in Florida. Phytopathology 98, 387–396 (2008).Article 
CAS 
PubMed 

Google Scholar 
Halbert, S. E. & Manjunath, K. L. Asian citrus psyllids (Sternorrhyncha: Psyllidae) and greening disease of citrus: A literature review and assessment of risk in Florida. Fla. Entomol. 87, 330–353 (2004).Article 

Google Scholar  More

Landscape use and anthropogenic influenceThe site could have had a specific and maybe extraordinary position in the microregion or in the trade networks41,42. The idea for creating trenches may have spread along trade routes—either as a habit of migrating people or as an ideology in the area of South and West Bohemia, Southern Germany, and the Austrian Land Salzburg55,56,57.Creeks along the settlement were major landscape elements. The settlement itself is entirely situated in the landscape periphery2. Steep slopes above Židova strouha creek and Blatenský potok brooks fundamentally limit agricultural use of the hinterland on the Březnice site, based on a model of reconstruction of the landscape potential (Fig. 6). The slopes may have been covered with sparse forest or shrubs. They were also forested in the nineteenth century, at the time of maximum agricultural load on the landscape as historical maps prove (Fig. 7).Figure 7Březnice and Hvožďany: the map of the second military mapping. Site catchements81 are according to the walk distance83 are shown hatched.Full size imageFieldsIn terms of human nutrition, the fields were crucial. The arable field area consisted of the actually cultivated fields and fallows. Analysis of plant macroremains provides us with knowledge of the grown species and the weed spectrum. The potential area and location of fields are reconstructed by a model that combines the agricultural potential of the landscape and previously published knowledge of the economic needs of the economic unit2,5,60,61,62,63.There is a possibility to assume, according to the SCA, the location of fields in relatively drier parts of the settlement area. Areas suitable for fields were probably located eastward and northward of the site, about 10–15 min walking distance (Fig. 6). The burial site was located beyond the northern border of the area where our analysis predicted the existence of fields93.Areas located eastward and northward of the settlement are even drier nowadays. The wetter fields may have been located in the north and northeast of the settlement, in its immediate vicinity. Moist soil is still present in these places today. The seeds and fruits of weed plants appear to have been transferred into the settlement together with the harvest. After being cleaned they were deposited as waste or used for further purposes, e.g. as an organic ingredient in ceramics or in daub4. The drier fields could correspond to finds of the following plant species: Arenaria serpyllifolia, Clinopodium acinos, Galeopsis augustifolia, Geranium cf. columbinum, Medicago lupulina, Rumex acetosella, Scleranthus annuus. Conversely, the following plants may have grown in the wetter fields, as documented in features on the settlement: Echinochloa crus-galli, Fumaria officinalis, Persicaria lapatifolia, Rumex cf. acetosa, Stachys arvensis.Synanthropic vegetation and ruderal habitatsArchaeobotanical analysis recorded many plant species characteristic for ruderal vegetation (most frequented Chenopodium album, Atriplex sp., Galium spurium, Polygonum aviculare, Chenopodium ficifolium, Fallopia convolvulus, Galium aparine). One could expect the presence of ruderals in the settlement area and its nearest surroundings in places that have been intensively used by humans and animals. The plants on the site could have reached the buildings by direct sedimentation and accidental charring, use of the ruderal plants, or as a result of waste burning.Deforested grazing areasGrazing took place in the enclosures and in the forests, which were made more open. The grazing of domestic animals had to be regulated in order to avoid crop damage and free movement around the settlement area. Winter fodder for animals had to be obtained within the reach of the settlement area, which contributed to the further lowering density of the forest. The archaeobotanical data reflect the grazing habitats in forest and deforested areas. Detrended correspondence analysis shows two clusters of plant species compatible with such environment (Fig. 4). The question is the process by which the plants reached the settlements. Species which appear in the ordinary space between the grassland and woodland—shrub positions could have grown on grasslands and light forests (e.g. Lychnis flos-cuculi, Dianthus cf. armeria, Galium palustre, Festuca ovina, Juncus sp., Campanula cf. glomerata) species in the ordinary space between “ruderal” and “grassland” could have grown at both habitats, e.g. at the transition of the settlement to the open countryside (e.g. Achillea millefolium, Alopecurus pratense, Asperula cynanchica, Briza media, Festuca cf. pratensis, Galium cf. verum, Ranunculus cf. bulbosus, Silene vulgaris, Stellaria graminea, Trifolium pratense). Taxa displayed between the “field” and “grassland” could have grown for example on fallow lands or abandoned fields that have successively overgrown (e.g. Clinopodinum acinus, Plantago lanceolata, Trifolium repens, Polycnemum arvense, Trifolium arvense). Taxa typical for “field” and “woodland-shrub” significantly differ in Březnice (Fig. 8).Figure 8Březnice: detrended correspondence analysis (DCA) Displayed samples and botanical taxa: the first axis explains 44.57% variability, the first and the second axis together 50.47%.Full size imageThe archaeobotanical analysis captured multiple grassland types. Both drier and wetter environments can be reconstructed. Wetter areas were represented by e.g. Alopecurus pratense, Alopecurus geniculatus, Carex cf. hirta, Carex cf. vulpina, cf. Euphorbia palustris, Galium cf. palustre, Juncus sp., Lychnis flos-cuculi, Myosotis sp., Persicaria lapatifolia, Plantago lanceolata, Stachys cf. palustris, Stellaria graminea, Urtica dioica. Drier areas were represented by e.g. Asperula cynanchica, Briza media, Campanula cf. glomerata, Carex cf. contigua, Clinopodium acinos, Dianthus cf. armeria, Phleum sp., Festuca cf. ovina, Galeopsis augustifolia, Galium cf. verum, Medicago lupulina, Polycnemum arvense, Ranunculus cf. bulbosus, Scleranthus annuus, Silene vulgaris, Solanum nigrum, Spergula arvensis, Trifolium arvense, Vicia tetrasperma, Vicia cf. villosa (Fig. 8).The existence of grasslands is associated with long-term human activities94. The Bechyně region has been apparently continuously settled since the end of the Early Bronze Age34. The landscape around the settlements has always been influenced by human activity and a large part of it has been deforested or covered with a sparse pastoral forest. However, not all the settlement areas were occupied permanently3, and those which were unoccupied became overgrown.Meadows and pastures are much more suitable for the grazing of herbivores than a forest with a dense canopy. Forest-steppe or significantly open forest is a convenient combination ensuring sufficient grazing for animals and wood production. Grazing increased soil fertility, reduced weeds on ruderal sites, and prevented forest growth95. Our study recorded a wide spectrum of charred macroremains of plants, which grew in the grasslands. They could have reached the site in several ways. In the excrements of the animals coming from a grazing area96, as raw materials collected by humans for further use in the settlement economy (e.g. food, medicinal plants, dyeing plants, bedding, admixture of screed and ceramic earth and daub, etc.). Studies1,3,32 assume, that the area in the immediate vicinity of the site was probably forestless. Forests at least half an hour’s walking distance from the site was significantly influenced by human activity. With an increasing distance from the centre of the site, the forest was probably less affected by human activities. The character of woodland usually clearly corresponded with the environmental conditions of the location31. The current forest area is extremely unsuitable for usage (slopes, wetlands). We assume that the occurrence of woodlands and shrubs in the Late Bronze Age was much more widespread, even in less extreme habitats.Shrubs and forestSpecies of herbs from different forest and shrub environments were also frequently recorded in the archaeobotanical assemblage. In the environment of wet forests could have grown e.g. Alliaria petiolata, Galium cf. palustre, Galium odoratum, Galium sylvaticum, Lychnis flos-cuculi, Persicaria lapatifolia, Solanum dulcamara, Stachys cf. palustris. In the coastal shrubs and edges of wet forests could have occured e.g. Cuscuta cf. europea, cf. Euphorbia palustris, Chelidinium majus, Impatiens nolitangere, Juncus sp., Myosoton aquaticum, Urtica dioica, Veronica hederifolia. Suitable locations could have been along the streams that flowed around the settlement and were within a quarter-hour walk. On the edges of the forests and their glades could have grown e.g. Atropa bella-donna, Festuca cf. ovina, Galium aparine, Prunella vulgaris, Rumex acetosella, Silene dioica, Thymus sp. Light forests and slopes were suitable for e.g. for Campanula cf. glomerata, Carex cf. contigua, Dianthus cf. armeria, Geranium cf. columbinum (Fig. 8).The areas for hunting and harvesting of wild crops were also economically important. The fruits that could have been collected included Corylus avellana, Crataegus sp., Atropa bella-donna, Prunus spinosa, Quercus sp., Rubus ideaus, Rubus fruticosus, Sambucus nigra, Solanum nigrum, Solanum dulcamara; their remains were found in the infills of features. The source of the collected fruits was located mostly in the sparse forest, forest edges and shrubs.The forest was also a source of building material and firewood3. From this acreage, the firewood for one farm could have been collected from 10 hectares. The rest would be used for collecting fodder and forest grazing7. The map of the potential natural vegetation92 predicts acidophilous oak forests (Quercetea robori-petraeae, Fig. 7) for the majority of both settlement areas. These species-poor woodlands are characteristic of Quercus dominance and in places mixed with Betula, Pinus, Sorbus, and Tilia on both dry and wet acidic soils, and Fagus, Abies, or Picea at higher altitudes. The results of our anthracological analysis clearly documented the predominance of this vegetation type in the vicinity of both archaeological sites.In the valleys of the streams and rivers were reconstructed alluvial forests with Alnus and mesophilous oak-hornbeam woods. The archeobotanical analysis of charcoals and fragments of fruits detected presence of Quercus, Tilia, Corylus, Crataegus, and Carpinus. These macroremains indicate existence of mesophilous forests. The hornbeam is rare in southern Bohemia97, it is the first of the archaeobotanical finds from prehistory. Due to the structure of taxa, which was captured by archaeobotanical analysis in Březnice, meadows and alder tree woods may be assumed there. Results of archaeobotanical analysis also documented the presence of Salix/Populus, Alnus.The most dominant tree species discovered in the trench-like features was oak which was mainly used as a construction material (Fig. 5). Firs were used as construction wood, which is predominantly present in stake pits in Březnice. In Hvožďany, trench 1 contained a cultural layer with apparent remains of a destructed building with charcoals of fir, spruce, and pine which in this case also served as construction wood34. The material commonly available in the forests surrounding the settlement area served as firewood (Figs. 4, 5, 8, 9).Figure 9Hvožďany: detrended correspondence analysis (DCA) Displayed samples and botanical taxa: the first axis explains 64.08% variability, the first and the second axis together 72.12%.Full size imageTime of housing: landscape potential vs. human needsThe homestead management (construction, abandonment, destruction, reconstruction etc.) during the settlement´s lifespan is a long-term studied question98,99. The existence of a hierarchized Late Bronze Age settlement network was evident in the lowland settlement areas of the Czech Republic with the continuity of occupational activity. Two main types of settlement are usually recognized there: (1) long-term large settlements and (2) short-term small settlements100,101. Agricultural productivity, exploitation of natural resources in settlements areas, and trade networks differed in cases of small or large settlements102. From the archaeological evidence perspective, the South Bohemia region was sparsely populated and the presence of long-term large settlements areas was very rare34.Previous research (excavations and magnetometry survey) has led to the conclusion that the 70 trenches are depositions of 70 houses and each trench is a deposition of one original house4,5,58. Based on such data, there could be many settlement forms differing in the space and time. The possible size of the settlement could be derived from the comparison of demands for fields, pastures, and forests with carrying capacity.SCA model and prediction model when compared to the possible demand7 of the community show that forest and pastures were not limiting factor for the settlement sustainability. In case of fields, there could be four variants of the possible extent of the settlement connected with different intensity of landuse. (1) The optimal acreage of fields (69 ha) with optimal land-use (7.5 ha/household); (2) the maximal extent of the fields 104 ha with optimal land-use or optimal extent of the field systems with intensive land-use (5 ha); (3) the maximal extent of the fields 104 ha and intensive land-use (5 ha); (4) sub-optimal land-use and fields located outside of the reach and optimal soils (Table 2). This model is an ideal prediction. For better yield the farmer could travel longer time than is expected however poor soils on a sloped terrain in the close vicinity were probably used rather as pastures.Table 2 Březnice: possible duration of the settlement based on four land use strategies: light green-optimal extent of the fields (69 hectares), with 7.5 hectares of fields per homestead; dark green-maximal extent of the fields (104 ha) or more intensive use of the fields (5 ha/homestead); maximal use and maximal extent; red—not sustainable agriculture or location of fields on places outside predicted optimal areas.Full size tableDrawing upon the typological and radiocarbon dating, it is often impossible to find out what was the lifespan of the settlement on the actual site. In this case, the uncertainty of 14C dates gives us a maximum possible span 73–264 years (95% probability), probably for 107–192 years (68% probability) (Supplementary Table 1, Fig. 2). Typological dating indicates 100–150 years (1150–1000 BC).The model described above indicates that the hinterland of Březnice could have sustained up to 20 houses at the same time in case of the maximal extent of the fields and intensive land-use. In this case, the settlement would have lasted only 90 years. If the land was used extensively it could have bore maximum of 14 houses at the time. That would correspond to a duration of roughly 126 years. Optimal areas of field systems in combination with sufficiently large fallows could have been used by a maximum of nine houses present at the time (192 years). The crucial part of the model is ritual burning and rebuilding houses after one generation58.Models of potential spatial and temporal characteristics of the settlement derived from prediction modeling cannot be tested. Therefore we need to compare our predictions with the radiocarbon model. The shortest duration of the settlement based on prediction is 90 years which corresponds with the 72 years modelled from 14C data. Since the model does not reflect the maximal duration of dwelling, this limit has to be based only on 14C model (262 years at 95% probability. At the maximum possible landuse levels, the settlement could have lasted from 72/90 to 262 years. The optimal duration of the settlement based on prediction could be 192–262 years. Extensive but more demanding land-use could support the duration of the settlement from 126 to 262 years (Table 2).Březnice and Hvožďany: the interpretation of both settlement areas from an archaeobotanical perspectiveThe two similarly dated settlement areas in one microregion with high quality archaeobotanical data allow (based on archaeobotanical material) a detailed study of the behaviour of communities in the Late Bronze Age. Archaeobotanical assemblages bring the reconstruction of the environment where the communities of the settlements drew plant resources from. Although the number of plant remains from both sites is significantly different, the interpretation of the environment does not differ in broad terms. For both sites, a similar share of fields and ruderals was documented. The spectrum of cultivated species was also identical41. Both settlements were self-sufficient in plant production—both waste and production parts of cultivated plants were found in the assemblages21,34,41. Animal bones were not preserved due to the acidic soil. However, for the Late Bronze Age sites the types of the domestic103 and the hunted104 animals are known.According to the environmental model, a greater proportion of species in Březnice came from grassland rather than from woodland and shrubs (Fig. 4). According to the analysis of plant macroremains more deforestation was recorded (i.e. more fields and pastures) in Březnice than in Hvožďany (Figs. 4, 5, 8, 9). Predicted areas for fields were in case of Hvožďany from 27 to 130 ha. Hvožďany site could possibly have larger field systems, but further away than in case of Březnice settlement. In Hvožďany there have been documented many taxa typical also for ruderal sites and fields. Several taxa could have grown either on ruderal sites or grasslands. Three reconstructed environments (ruderals, fields, grasslands) in Hvožďany significantly differ from woodland—shrub (Figs. 8, 9). The large volume of analysed samples from Březnice brought a number of botanical taxa which was mostly found in only a few specimens but ultimately brought the opportunity to reconstruct the surroundings of the site in more detail. In Hvožďany, a common spectrum of plants was found (Fig. 9), which usually occurs at similar South Bohemian sites, e.g. Černýšovice, Rataje, Zhoř, Oldřichov, Písek—Bakaláře105,106. Nevertheless, it brings the possibility to reconstruct the surroundings at least in rough features.The archaeological field data does not allow us to reconstruct how many houses were on the Hvožďany site at the same time. Total inhabited area of ​​the settlement in Březnice is approximately 13 ha, at Hvožďany site it is altogether 5 ha. It suggests two explanations: either more people lived in Březnice than in Hvožďany or the settlement had a longer span (or both possibilities). However, both options mean greater deforestation in Březnice. The carrying capacity and landscape potential of the settlement in Hvožďany could not have been exhausted (Fig. 6). The area of high quality soil in a quarter/half hour’s walk from the site is sufficient for 3.6–25 houses (27–130 ha). Two community areas could have been separated by the Lužnice river (walking distance within one hour). The agricultural systems of the settlements were probably very similar. According to our models, both settlement sites would have only needed to exploit natural resources in their immediate hinterland, within an hour walking radius. The limiting factor is the availability of suitable land for fields.According to the archaeobotanical results, the landscape in Březnice was more affected by human activity than the one in Hvožďany. A greater number of species were found, evidenced by light woodland and shrubs and different types of grassland. In the vicinity of the settlement from which people drew resources, a light landscape can be assumed. So far there is no pollen profile available. Approximately 2 m of accumulated clay and sand without organics were sampled in the floodplain of the Židova strouha. About 20 km away from Březnice, the analysis was performed in Sepekov, which base could have corresponded to the Bronze Age (2920 ± 410 BP). The character of the vegetation based on the profile could be interpreted as wet and relatively nutritious fir woodland or fir alder woodland situated on a relatively small spring area at the edge of the water meadow of the Smutná river. The palaeobotanical record in this phase does not record any effect of the settlement on the vegetation present34. The profile containing the pollen record from the Borkovická blata is located about 10 km away from Březnice. As well as the profile from Sepekov, it reflects local peat bog vegetation of the subboreal character without significant indicators of human activity107.The conditions and availability of resources in the hinterland of both settlements were probably overall so good that the details did not matter much. In the vicinity of both settlements, there were a sufficient number of areas for fields, pastures, and cultural forests. The settlement areas of the Late Bronze Age in South Bohemia were probably in separate deforested niches. More

Wikelski, M. & Cooke, S. J. Conservation physiology. Trends Ecol. Evol. 21, 38–46 (2006).Article 
PubMed 

Google Scholar 
Cattet, M. et al. Understanding grizzly bear health in the context of changing landscapes. Foothills Model Forest Grizzly Bear Research Program Annual Report, 80–86 (2005).Reeder, D. M. & Kramer, K. M. Stress in free-ranging mammals: Integrating physiology, ecology, and natural history. J. Mammal. 86, 225–235. https://doi.org/10.1644/bhe-003.1 (2005).Article 

Google Scholar 
Dunlop, R. A., Braithwaite, J., Mortensen, L. O. & Harris, C. M. Assessing population-level effects of anthropogenic disturbance on a marine mammal population. Front. Mar. Sci. 8, 230 (2021).Article 

Google Scholar 
Atkinson, S., Crocker, D., Houser, D. & Mashburn, K. Stress physiology in marine mammals: How well do they fit the terrestrial model?. J. Comp. Physiol. B. 185, 463–486. https://doi.org/10.1007/s00360-015-0901-0 (2015).Article 
CAS 
PubMed 

Google Scholar 
Romero, L. M. & Beattie, U. K. Common myths of glucocorticoid function in ecology and conservation. J. Exp. Zool. Part A: Ecol. Integr. Physiol. 337, 7–14. https://doi.org/10.1002/jez.2459 (2022).Article 
CAS 

Google Scholar 
Champagne, C. D. et al. Blubber cortisol qualitatively reflects circulating cortisol concentrations in bottlenose dolphins. Mar. Mamm. Sci. 33, 134–153 (2017).Article 
CAS 

Google Scholar 
Champagne, C. D. et al. Comprehensive endocrine response to acute stress in the bottlenose dolphin from serum, blubber, and feces. Gen. Comp. Endocrinol. 266, 178 (2018).Article 
CAS 
PubMed 

Google Scholar 
Teerlink, S., Horstmann, L. & Witteveen, B. Humpback whale (Megaptera novaeangliae) blubber steroid hormone concentration to evaluate chronic stress response from whale-watching vessels. Aquat. Mamm. 44, 411 (2018).Article 

Google Scholar 
Mingramm, F. M., Keeley, T., Whitworth, D. J. & Dunlop, R. A. Blubber cortisol levels in humpback whales (Megaptera novaeangliae): A measure of physiological stress without effects from sampling. Gen. Comp. Endocrinol. 291, 113436 (2020).Article 
CAS 
PubMed 

Google Scholar 
Saco, Y. et al. Evaluation of serum cortisol, metabolic parameters, acute phase proteins and faecal corticosterone as indicators of stress in cows. Vet. J. 177, 439–441 (2008).Article 
CAS 
PubMed 

Google Scholar 
Rolland, R. M. et al. Evidence that ship noise increases stress in right whales. Proc. R. Soc. B: Biol. Sci. 279, 2363–2368. https://doi.org/10.1098/rspb.2011.2429 (2012).Article 

Google Scholar 
Rocha, R., Clapham, P. J. & Ivashchenko, Y. V. Emptying the oceans: A summary of industrial whaling catches in the 20th century. Mar. Fish. Rev 76, 37–48 (2014).Article 

Google Scholar 
Comission, I. W. Report of the scientific committee. Journal of Cetacean Research and Management SC/68C (2021).Ducklow, H. W. et al. West Antarctic Peninsula: An ice-dependent coastal marine ecosystem in transition. Oceanography 26, 190–203 (2013).Article 

Google Scholar 
Laws, R. Seals and whales of the Southern Ocean 81–96 (Philosophical Transactions of the Royal Society of London. Series B, 1977).
Google Scholar 
Savoca, M. S. et al. Baleen whale prey consumption based on high-resolution foraging measurements. Nature 599, 85–90. https://doi.org/10.1038/s41586-021-03991-5 (2021).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Pallin, L. J. et al. High pregnancy rates in humpback whales (Megaptera novaeangliae) around the Western Antarctic Peninsula, evidence of a rapidly growing population. R. Soc. Open Sci. https://doi.org/10.1098/rsos.180017 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Bender, N. A., Crosbie, K. & Lynch, H. J. Patterns of tourism in the Antarctic Peninsula region: A 20-year analysis. Antarct. Sci. 28, 194–203. https://doi.org/10.1017/s0954102016000031 (2016).Article 
ADS 

Google Scholar 
Operators, I. A. o. A. T. (2019).Trumble, S. J. et al. Baleen whale cortisol levels reveal a physiological response to 20th century whaling. Nat. Commun. https://doi.org/10.1038/s41467-018-07044-w (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Cates, K. A. et al. Corticosterone in central north pacific male humpback whales (Megaptera novaeangliae): Pairing sighting histories with endocrine markers to assess stress. Gen. Comp. Endocrinol. 296, 113540 (2020).Article 
CAS 
PubMed 

Google Scholar 
Sprogis, K. R., Videsen, S. & Madsen, P. T. Vessel noise levels drive behavioural responses of humpback whales with implications for whale-watching. eLife https://doi.org/10.7554/elife.56760 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Rolland, R. et al. Fecal glucocorticoids and anthropogenic injury and mortality in North Atlantic right whales Eubalaena glacialis. Endanger. Species Res. 34, 417–429. https://doi.org/10.3354/esr00866 (2017).Article 

Google Scholar 
Modest, M. et al. First description of migratory behavior of humpback whales from an Antarctic feeding ground to a tropical calving ground. Anim. Biotelem. 9, 1–16 (2021).Article 

Google Scholar 
Amaral, R. S. Use of alternative matrices to monitor steroid hormones in aquatic mammals: A review. Aquat. Mamm. 36, 162 (2010).Article 

Google Scholar 
Graham, K. M., Burgess, E. A. & Rolland, R. M. Stress and reproductive events detected in North Atlantic right whale blubber using a simplified hormone extraction protocol. Conserv. Physiol. https://doi.org/10.1093/conphys/coaa133 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Atkinson, S. et al. Pregnancy rate and biomarker validations from the blubber of eastern North Pacific blue whales. Mar. Mamm. Sci. https://doi.org/10.1111/mms.12616 (2019).Article 

Google Scholar 
Hunt, K. E., Rolland, R. M., Kraus, S. D. & Wasser, S. K. Analysis of fecal glucocorticoids in the North Atlantic right whale (Eubalaena glacialis). Gen. Comp. Endocrinol. 148, 260–272. https://doi.org/10.1016/j.ygcen.2006.03.012 (2006).Article 
CAS 
PubMed 

Google Scholar 
Mashburn, K. L. & Atkinson, S. Evaluation of adrenal function in serum and feces of Steller sea lions (Eumetopias jubatus): Influences of molt, gender, sample storage, and age on glucocorticoid metabolism. Gen. Comp. Endocrinol. 136, 371–381. https://doi.org/10.1016/j.ygcen.2004.01.016 (2004).Article 
CAS 
PubMed 

Google Scholar 
Jeanniard Du Dot, T. et al. Changes in glucocorticoids, IGF-I and thyroid hormones as indicators of nutritional stress and subsequent refeeding in Steller sea lions (Eumetopias jubatus). Comp. Biochem. Physiol. Part A Mol. Integr. Physiol 152, 524–534. https://doi.org/10.1016/j.cbpa.2008.12.010 (2009).Article 
CAS 

Google Scholar 
Foley, C. A. H., Papageorge, S. & Wasser, S. K. Noninvasive stress and reproductive measures of social and ecological pressures in free-ranging African elephants. Conserv. Biol. 15, 1134–1142. https://doi.org/10.1046/j.1523-1739.2001.0150041134.x (2001).Article 

Google Scholar 
Challis, J. R., Matthews, S. G., Gibb, W. & Lye, S. J. Endocrine and paracrine regulation of birth at term and preterm. Endocr. Rev. 21, 514–550 (2000).CAS 

Google Scholar 
Robeck, T. R., Steinman, K. J. & O’Brien, J. K. Characterization and longitudinal monitoring of serum androgens and glucocorticoids during normal pregnancy in the killer whale (Orcinus orca). Gen. Comp. Endocrinol. 247, 116–129 (2017).Article 
CAS 
PubMed 

Google Scholar 
Rolland, R. M., Hunt, K. E., Kraus, S. D. & Wasser, S. K. Assessing reproductive status of right whales (Eubalaena glacialis) using fecal hormone metabolites. Gen. Comp. Endocrinol. 142, 308–317 (2005).Article 
CAS 
PubMed 

Google Scholar 
Burgess, E. A., Hunt, K. E., Kraus, S. D. & Rolland, R. M. Adrenal responses of large whales: Integrating fecal aldosterone as a complementary biomarker to glucocorticoids. Gen. Comp. Endocrinol. 252, 103–110. https://doi.org/10.1016/j.ygcen.2017.07.026 (2017).Article 
CAS 
PubMed 

Google Scholar 
Ducklow, H. W. et al. Marine pelagic ecosystems: The west Antarctic Peninsula. Philos. Trans. R. Soc. London B: Biol. Sci. 362, 67–94 (2007).Article 
PubMed 

Google Scholar 
Rogers, A. et al. Antarctic futures: An assessment of climate-driven changes in ecosystem structure, function, and service provisioning in the Southern Ocean. Ann. Rev. Mar. Sci. 12, 87–120 (2020).Article 
CAS 
PubMed 

Google Scholar 
Hillbrand, F. & Elsaesser, F. Concentrations of progesterone in the backfat of pigs during the oestrous cycle and after ovariectomy. J. Reprod. Fertil. 69, 73–80 (1983).Article 
CAS 
PubMed 

Google Scholar 
Funasaka, N. et al. Seasonal difference of diurnal variations in serum melatonin, cortisol, testosterone, and rectal temperature in Indo-Pacific bottlenose dolphins (Tursiops aduncus). Aquat. Mamm. 37, 433 (2011).Article 

Google Scholar 
Oki, C. & Atkinson, S. Diurnal patterns of cortisol and thyroid hormones in the Harbor seal (Phoca vitulina) during summer and winter seasons. Gen. Comp. Endocrinol. 136, 289–297. https://doi.org/10.1016/j.ygcen.2004.01.007 (2004).Article 
CAS 
PubMed 

Google Scholar 
Lavigne, D., Innes, S., Worthy, G. & Edwards, E. F. Lower critical temperatures of blue whales, Balaenoptera musculus. J. Theor. Biol. 144, 249–257 (1990).Article 
ADS 

Google Scholar 
Nichols, R. C. et al. Intra-seasonal variation in feeding rates and diel foraging behaviour in a seasonally fasting mammal, the humpback whale. R. Soc. Open Sci. https://doi.org/10.1098/rsos.211674 (2022).Article 
PubMed 
PubMed Central 

Google Scholar 
Bierlich, K. C. Incorporating Photogrammetric Uncertainty in UAS-based Morphometric Measurements of Baleen Whales, (2021).Kellar, N. M. et al. Blubber cortisol: A potential tool for assessing stress response in free-ranging dolphins without effects due to sampling. PLoS ONE 10, e0115257 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Bejder, L. & Samuels, A. Evaluating the effects of nature-based tourism on cetaceans. Mar. Mamm. Fish. Tour. Manag. Issues 1, 229–256 (2003).
Google Scholar 
New, L. F. et al. The modelling and assessment of whale-watching impacts. Ocean Coast. Manag. 115, 10–16 (2015).Article 

Google Scholar 
Avila, I. C., Correa, L. M. & Parsons, E. Whale-watching activity in Bahía Málaga, on the Pacific coast of Colombia, and its effect on humpback whale (Megaptera novaeangliae) behavior. Tour. Mar. Environ. 11, 19–32 (2015).Article 

Google Scholar 
Amrein, A. M., Guzman, H. M., Surrey, K. C., Polidoro, B. & Gerber, L. R. Impacts of whale watching on the behavior of humpback whales (Megaptera novaeangliae) in the Coast of Panama. Front. Mar. Sci. 7, 1105 (2020).Article 

Google Scholar 
Heenehan, H. et al. Caribbean Sea soundscapes: monitoring humpback whales, biological sounds, geological events, and anthropogenic impacts of vessel noise. Front. Mar. Sci., 347 (2019).Keay, J. M., Singh, J., Gaunt, M. C. & Kaur, T. Fecal glucocorticoids and their metabolites as indicators of stress in various mammalian species: A literature review. J. Zoo Wildl. Med. 37, 234–244 (2006).Article 
PubMed 

Google Scholar 
Harris, K., Gende, S. M., Logsdon, M. G. & Klinger, T. Spatial pattern analysis of cruise ship-humpback whale interactions in and near Glacier Bay National Park, Alaska. Environ. Manag. 49, 44–54. https://doi.org/10.1007/s00267-011-9754-9 (2012).Article 
ADS 

Google Scholar 
Palsbøll, P. J., Larsen, F. & Hansen, E. S. Sampling of skin biopsies from free-raging large cetaceans in west greenland: Development of new biopsy tips and bolt designs. International Whaling Commission Special Issue Series (1991).Weinstein, B. G., Double, M., Gales, N., Johnston, D. W. & Friedlaender, A. S. Identifying overlap between humpback whale foraging grounds and the Antarctic krill fishery. Biol. Cons. 210, 184–191 (2017).Article 

Google Scholar 
Lambertsen, R. H. A biopsy system for large whales and its use for cytogenetics. J. Mammal. 68, 443–445. https://doi.org/10.2307/1381495 (1987).Article 

Google Scholar 
Gilson, A., Syvanen, M., Levine, K. & Banks, J. Deer gender determination by polymerase chain reaction. Calif. Fish Game 84, 159–169 (1998).
Google Scholar 
Aasen, E. & Medrano, J. F. Amplification of the ZFY and ZFX genes for sex identification in humans, cattle, sheep and goats. Bio/Technology 8, 1279–1281 (1990).CAS 
PubMed 

Google Scholar 
Valsecchi, E. & Amos, W. Microsatellite markers for the study of cetacean populations. Mol. Ecol. 5, 151–156 (1996).Article 
CAS 
PubMed 

Google Scholar 
Palsbøll, P., Bérubé, M., Larsen, A. & Jørgensen, H. Primers for the amplification of tri-and tetramer microsatellite loci in baleen whales. Mol. Ecol. 6, 893–895 (1997).Article 
PubMed 

Google Scholar 
Berube, M., Jørgensen, H., McEwing, R. & Palsbøll, P. J. Polymorphic di-nucleotide microsatellite loci isolated from the humpback whale, Megaptera novaeangliae. Mol. Ecol. 9, 2181–2183 (2000).Article 
CAS 
PubMed 

Google Scholar 
Waldick, R., Brown, M. & White, B. Characterization and isolation of microsatellite loci from the endangered North Atlantic right whale. Mol. Ecol. 8, 1763–1765 (1999).Article 
CAS 
PubMed 

Google Scholar 
Baker, C. S. et al. Strong maternal fidelity and natal philopatry shape genetic structure in North Pacific humpback whales. Mar. Ecol. Progress Ser. 494, 291 (2013).Article 
ADS 

Google Scholar 
Constantine, R. et al. Abundance of humpback whales in Oceania using photo-identification and microsatellite genotyping. Mar. Ecol. Prog. Ser. 453, 249–261 (2012).Article 
ADS 

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

Google Scholar 
Kalinowski, S. T., Taper, M. L. & Marshall, T. C. Revising how the computer program CERVUS accommodates genotyping error increases success in paternity assignment. Mol. Ecol. 16, 1099–1106 (2007).Article 
PubMed 

Google Scholar 
Kellar, N. M., Trego, M. L., Marks, C. I. & Dizon, A. E. Determining pregnancy from blubber in three species of delphinids. Mar. Mamm. Sci. 22, 1–16 (2006).Article 

Google Scholar 
Pallin, L., Robbins, J., Kellar, N., Bérubé, M. & Friedlaender, A. Validation of a blubber-based endocrine pregnancy test for humpback whales. Conserv. Physiol. https://doi.org/10.1093/conphys/coy031 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Kellar, N. M. et al. Low reproductive success rates of common bottlenose dolphins Tursiops truncatus in the northern Gulf of Mexico following the Deepwater Horizon disaster (2010–2015). Endanger. Species Res. 33, 143–158 (2017).Article 

Google Scholar 
R: A language and environment for statistical computing. (R Foundation for Statistical Computing, Vienna, Austria, 2021).ArcMap (version 10.8.2) (Redlands, CA: Esri Inc, 2022). More

Annual grains, domesticated from wild species, have dominated agriculture since the Neolithic. A new study reports how turning to high-yield perennial rice crops could maintain key ecosystem functions while supporting livelihoods.The past several decades have seen modest but growing investments in the development of perennial grain crops, including perennial counterparts of wheat, rice and sorghum suitable for the USA, China, Europe and Africa. One technique involves domesticating wild perennial species through continual selection of desirable traits over multiple generations3. A recently developed perennial grain currently grown for niche markets in the USA, Kernza, was domesticated from Thinopyrum intermedium, a wild relative of wheat. While yields of Kernza remain low compared with those of annual wheat, they are increasing. As with the development of perennial rice, plant breeders can also cross perennial species with domesticated annual relatives to produce perennial hybrids with desirable traits derived from the annual parent3. More

.readcube-buybox { display: none !important;}
More than 30% of shark and ray species are edging towards extinction, mainly because they are unintentionally caught by fishers targeting tuna and other commercially valuable species. A new device might help to keep some of these threatened species away from fishing hooks.

Access options

/* style specs start */
style{display:none!important}.LiveAreaSection-193358632 *{align-content:stretch;align-items:stretch;align-self:auto;animation-delay:0s;animation-direction:normal;animation-duration:0s;animation-fill-mode:none;animation-iteration-count:1;animation-name:none;animation-play-state:running;animation-timing-function:ease;azimuth:center;backface-visibility:visible;background-attachment:scroll;background-blend-mode:normal;background-clip:borderBox;background-color:transparent;background-image:none;background-origin:paddingBox;background-position:0 0;background-repeat:repeat;background-size:auto auto;block-size:auto;border-block-end-color:currentcolor;border-block-end-style:none;border-block-end-width:medium;border-block-start-color:currentcolor;border-block-start-style:none;border-block-start-width:medium;border-bottom-color:currentcolor;border-bottom-left-radius:0;border-bottom-right-radius:0;border-bottom-style:none;border-bottom-width:medium;border-collapse:separate;border-image-outset:0s;border-image-repeat:stretch;border-image-slice:100%;border-image-source:none;border-image-width:1;border-inline-end-color:currentcolor;border-inline-end-style:none;border-inline-end-width:medium;border-inline-start-color:currentcolor;border-inline-start-style:none;border-inline-start-width:medium;border-left-color:currentcolor;border-left-style:none;border-left-width:medium;border-right-color:currentcolor;border-right-style:none;border-right-width:medium;border-spacing:0;border-top-color:currentcolor;border-top-left-radius:0;border-top-right-radius:0;border-top-style:none;border-top-width:medium;bottom:auto;box-decoration-break:slice;box-shadow:none;box-sizing:border-box;break-after:auto;break-before:auto;break-inside:auto;caption-side:top;caret-color:auto;clear:none;clip:auto;clip-path:none;color:initial;column-count:auto;column-fill:balance;column-gap:normal;column-rule-color:currentcolor;column-rule-style:none;column-rule-width:medium;column-span:none;column-width:auto;content:normal;counter-increment:none;counter-reset:none;cursor:auto;display:inline;empty-cells:show;filter:none;flex-basis:auto;flex-direction:row;flex-grow:0;flex-shrink:1;flex-wrap:nowrap;float:none;font-family:initial;font-feature-settings:normal;font-kerning:auto;font-language-override:normal;font-size:medium;font-size-adjust:none;font-stretch:normal;font-style:normal;font-synthesis:weight style;font-variant:normal;font-variant-alternates:normal;font-variant-caps:normal;font-variant-east-asian:normal;font-variant-ligatures:normal;font-variant-numeric:normal;font-variant-position:normal;font-weight:400;grid-auto-columns:auto;grid-auto-flow:row;grid-auto-rows:auto;grid-column-end:auto;grid-column-gap:0;grid-column-start:auto;grid-row-end:auto;grid-row-gap:0;grid-row-start:auto;grid-template-areas:none;grid-template-columns:none;grid-template-rows:none;height:auto;hyphens:manual;image-orientation:0deg;image-rendering:auto;image-resolution:1dppx;ime-mode:auto;inline-size:auto;isolation:auto;justify-content:flexStart;left:auto;letter-spacing:normal;line-break:auto;line-height:normal;list-style-image:none;list-style-position:outside;list-style-type:disc;margin-block-end:0;margin-block-start:0;margin-bottom:0;margin-inline-end:0;margin-inline-start:0;margin-left:0;margin-right:0;margin-top:0;mask-clip:borderBox;mask-composite:add;mask-image:none;mask-mode:matchSource;mask-origin:borderBox;mask-position:0 0;mask-repeat:repeat;mask-size:auto;mask-type:luminance;max-height:none;max-width:none;min-block-size:0;min-height:0;min-inline-size:0;min-width:0;mix-blend-mode:normal;object-fit:fill;object-position:50% 50%;offset-block-end:auto;offset-block-start:auto;offset-inline-end:auto;offset-inline-start:auto;opacity:1;order:0;orphans:2;outline-color:initial;outline-offset:0;outline-style:none;outline-width:medium;overflow:visible;overflow-wrap:normal;overflow-x:visible;overflow-y:visible;padding-block-end:0;padding-block-start:0;padding-bottom:0;padding-inline-end:0;padding-inline-start:0;padding-left:0;padding-right:0;padding-top:0;page-break-after:auto;page-break-before:auto;page-break-inside:auto;perspective:none;perspective-origin:50% 50%;pointer-events:auto;position:static;quotes:initial;resize:none;right:auto;ruby-align:spaceAround;ruby-merge:separate;ruby-position:over;scroll-behavior:auto;scroll-snap-coordinate:none;scroll-snap-destination:0 0;scroll-snap-points-x:none;scroll-snap-points-y:none;scroll-snap-type:none;shape-image-threshold:0;shape-margin:0;shape-outside:none;tab-size:8;table-layout:auto;text-align:initial;text-align-last:auto;text-combine-upright:none;text-decoration-color:currentcolor;text-decoration-line:none;text-decoration-style:solid;text-emphasis-color:currentcolor;text-emphasis-position:over right;text-emphasis-style:none;text-indent:0;text-justify:auto;text-orientation:mixed;text-overflow:clip;text-rendering:auto;text-shadow:none;text-transform:none;text-underline-position:auto;top:auto;touch-action:auto;transform:none;transform-box:borderBox;transform-origin:50% 50%0;transform-style:flat;transition-delay:0s;transition-duration:0s;transition-property:all;transition-timing-function:ease;vertical-align:baseline;visibility:visible;white-space:normal;widows:2;width:auto;will-change:auto;word-break:normal;word-spacing:normal;word-wrap:normal;writing-mode:horizontalTb;z-index:auto;-webkit-appearance:none;-moz-appearance:none;-ms-appearance:none;appearance:none;margin:0}.LiveAreaSection-193358632{width:100%}.LiveAreaSection-193358632 .login-option-buybox{display:block;width:100%;font-size:17px;line-height:30px;color:#222;padding-top:30px;font-family:Harding,Palatino,serif}.LiveAreaSection-193358632 .additional-access-options{display:block;font-weight:700;font-size:17px;line-height:30px;color:#222;font-family:Harding,Palatino,serif}.LiveAreaSection-193358632 .additional-login >li:not(:first-child)::before{transform:translateY(-50%);content:””;height:1rem;position:absolute;top:50%;left:0;border-left:2px solid #999}.LiveAreaSection-193358632 .additional-login >li:not(:first-child){padding-left:10px}.LiveAreaSection-193358632 .additional-login >li{display:inline-block;position:relative;vertical-align:middle;padding-right:10px}.BuyBoxSection-683559780{display:flex;flex-wrap:wrap;flex:1;flex-direction:row-reverse;margin:-30px -15px 0}.BuyBoxSection-683559780 .box-inner{width:100%;height:100%}.BuyBoxSection-683559780 .readcube-buybox{background-color:#f3f3f3;flex-shrink:1;flex-grow:1;flex-basis:255px;background-clip:content-box;padding:0 15px;margin-top:30px}.BuyBoxSection-683559780 .subscribe-buybox{background-color:#f3f3f3;flex-shrink:1;flex-grow:4;flex-basis:300px;background-clip:content-box;padding:0 15px;margin-top:30px}.BuyBoxSection-683559780 .subscribe-buybox-nature-plus{background-color:#f3f3f3;flex-shrink:1;flex-grow:4;flex-basis:100%;background-clip:content-box;padding:0 15px;margin-top:30px}.BuyBoxSection-683559780 .title-readcube{display:block;margin:0;margin-right:20%;margin-left:20%;font-size:24px;line-height:32px;color:#222;padding-top:30px;text-align:center;font-family:Harding,Palatino,serif}.BuyBoxSection-683559780 .title-buybox{display:block;margin:0;margin-right:29%;margin-left:29%;font-size:24px;line-height:32px;color:#222;padding-top:30px;text-align:center;font-family:Harding,Palatino,serif}.BuyBoxSection-683559780 .title-asia-buybox{display:block;margin:0;margin-right:5%;margin-left:5%;font-size:24px;line-height:32px;color:#222;padding-top:30px;text-align:center;font-family:Harding,Palatino,serif}.BuyBoxSection-683559780 .asia-link{color:#069;cursor:pointer;text-decoration:none;font-size:1.05em;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:1.05em6}.BuyBoxSection-683559780 .access-readcube{display:block;margin:0;margin-right:10%;margin-left:10%;font-size:14px;color:#222;padding-top:10px;text-align:center;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:20px}.BuyBoxSection-683559780 .access-asia-buybox{display:block;margin:0;margin-right:5%;margin-left:5%;font-size:14px;color:#222;padding-top:10px;text-align:center;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:20px}.BuyBoxSection-683559780 .access-buybox{display:block;margin:0;margin-right:30%;margin-left:30%;font-size:14px;color:#222;opacity:.8px;padding-top:10px;text-align:center;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:20px}.BuyBoxSection-683559780 .usps-buybox{display:block;margin:0;margin-right:30%;margin-left:30%;font-size:14px;color:#222;opacity:.8px;text-align:center;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:20px}.BuyBoxSection-683559780 .price-buybox{display:block;font-size:30px;color:#222;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;padding-top:30px;text-align:center}.BuyBoxSection-683559780 .price-from{font-size:14px;padding-right:10px;color:#222;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:20px}.BuyBoxSection-683559780 .issue-buybox{display:block;font-size:13px;text-align:center;color:#222;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:19px}.BuyBoxSection-683559780 .no-price-buybox{display:block;font-size:13px;line-height:18px;text-align:center;padding-right:10%;padding-left:10%;padding-bottom:20px;padding-top:30px;color:#222;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif}.BuyBoxSection-683559780 .vat-buybox{display:block;margin-top:5px;margin-right:20%;margin-left:20%;font-size:11px;color:#222;padding-top:10px;padding-bottom:15px;text-align:center;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;line-height:17px}.BuyBoxSection-683559780 .button-container{display:flex;padding-right:20px;padding-left:20px;justify-content:center}.BuyBoxSection-683559780 .button-container >*{flex:1px}.BuyBoxSection-683559780 .button-container >a:hover,.Button-505204839:hover,.Button-1078489254:hover,.Button-2808614501:hover{text-decoration:none}.BuyBoxSection-683559780 .readcube-button{background:#fff;margin-top:30px}.BuyBoxSection-683559780 .button-asia{background:#069;border:1px solid #069;border-radius:0;cursor:pointer;display:block;padding:9px;outline:0;text-align:center;text-decoration:none;min-width:80px;margin-top:75px}.BuyBoxSection-683559780 .button-label-asia,.ButtonLabel-3869432492,.ButtonLabel-3296148077,.ButtonLabel-1566022830{display:block;color:#fff;font-size:17px;line-height:20px;font-family:-apple-system,BlinkMacSystemFont,”Segoe UI”,Roboto,Oxygen-Sans,Ubuntu,Cantarell,”Helvetica Neue”,sans-serif;text-align:center;text-decoration:none;cursor:pointer}.Button-505204839,.Button-1078489254,.Button-2808614501{background:#069;border:1px solid #069;border-radius:0;cursor:pointer;display:block;padding:9px;outline:0;text-align:center;text-decoration:none;min-width:80px;max-width:320px;margin-top:10px}.Button-505204839 .readcube-label,.Button-1078489254 .readcube-label,.Button-2808614501 .readcube-label{color:#069}
/* style specs end */Subscribe to Nature+Get immediate online access to Nature and 55 other Nature journal$29.99monthlySubscribe to JournalGet full journal access for 1 year$199.00only $3.90 per issueAll prices are NET prices.VAT will be added later in the checkout.Tax calculation will be finalised during checkout.Buy articleGet time limited or full article access on ReadCube.$32.00All prices are NET prices.

Additional access options:

doi: https://doi.org/10.1038/d41586-022-03776-4

References

Subjects

Conservation biology More