Ecology

Mack, R. N. et al. Biotic invasions: causes, epidemiology, global consequences, and control. Ecol. Appl. 10, 689–710. https://doi.org/10.1890/1051-0761 (2000).Article 

Google Scholar 
Dukes, J. S. & Mooney, H. A. Disruption of ecosystem processes in western North America by invasive species. Rev. Chil. Hist. Nat. 77, 411–437 (2004).Article 

Google Scholar 
Vitousek, P. M. Biological invasions and ecosystem processes: towards an integration of population biology and ecosystem studies. Oikos 57, 7–13. https://doi.org/10.2307/3565731 (1990).Article 

Google Scholar 
Richardson, D. M. et al. Naturalization and invasion of alien plants: concepts and definitions. Diver. Distrib. 6, 93–107 (2000).Article 

Google Scholar 
Theoharides, K. A. & Dukes, J. S. Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion. New Phytol. 176, 256–273 (2007).Article 

Google Scholar 
Pyšek, P. et al. Naturalization of central European plants in North America: species traits, habitats, propagule pressure, residence time. Ecology 96, 762–774. https://doi.org/10.1890/14-1005.1 (2015).Article 
PubMed 

Google Scholar 
Estrada, J. A., Wilson, C. H. & Flory, S. L. Clonal integration enhances performance of an invasive grass. Oikos https://doi.org/10.1111/oik.07016 (2020).Article 

Google Scholar 
Otfinowski, R. & Kenkel, N. C. Clonal integration facilitates the proliferation of smooth brome clones invading northern fescue prairies. Plant Ecol. 199, 235–242. https://doi.org/10.1007/s11258-008-9428-8 (2008).Article 

Google Scholar 
Pyšek, P. & Richardson, D. M. in Biological Invasions (ed N. Nentwig) pp. 97–125 (Springer, New York, 2007).Klimešová, J. & Klimeš, L. Clonal growth diversity and bud banks of plants in the Czech flora: an evaluation using the CLO-PLA3 database. Preslia 80, 255–275 (2008).
Google Scholar 
Klimešová, J. et al. Handbook of standardized protocols for collecting plant modularity traits. Persp. Plant Ecol. https://doi.org/10.1016/j.ppees.2019.125485 (2019).Article 

Google Scholar 
Wang, Y. J. et al. Invasive alien plants benefit more from clonal integration in heterogeneous environments than natives. New Phytol. 216, 1072–1078 (2017).Article 

Google Scholar 
Klimešová, J. in Encyclopedia of Invasive Introduced Species (eds D. Simberloff & M. Reimanek) pp. 678–679 (University of California Press, California, 2011).Ott, J. P., Klimešová, J. & Hartnett, D. C. The ecology and significance of below-ground bud banks in plants. Ann. Bot. Lond. 123, 1099–1118. https://doi.org/10.1093/aob/mcz051 (2019).Article 

Google Scholar 
Sanchez, J. M., Sanchez, C. & Navarro, L. Can asexual reproduction by plant fragments help to understand the invasion of the NW Iberian coast by Spartina patens? Flora 257, 151410. https://doi.org/10.1016/j.flora.2019.05.009 (2019).Speek, T. A. A. et al. Factors relating to regional and local success of exotic plant species in their new range. Diver. Distrib. 17, 542–551 (2011).Article 

Google Scholar 
Wang, J. Y. et al. A meta-analysis of effects of physiological integration in clonal plants under homogeneous vs heterogeneous environments. Funct. Ecol. https://doi.org/10.1111/1365-2435.13732 (2020).Article 

Google Scholar 
Maurer, D. A. & Zedler, J. B. Differential invasion of a wetland grass explained by tests of nutrients and light availability on establishment and clonal growth. Oecologia 131, 279–288. https://doi.org/10.1007/s00442-002-0886-8 (2002).ADS 
Article 
PubMed 

Google Scholar 
Mueller, I. M. & Weaver, J. E. Relative drought resistance of seedlings of dominant prairie grasses. Ecology 23, 387–398 (1942).Article 

Google Scholar 
Vetter, V. M. S. et al. Invasion windows for a global legume invader are revealed after joint examination of abiotic and biotic filters. Plant Biol. 21, 832–843. https://doi.org/10.1111/plb.12987 (2019).CAS 
Article 
PubMed 

Google Scholar 
Ibanez, I. et al. Integrated assessment of biological invasions. Ecol. Appl. 24, 25–37. https://doi.org/10.1890/13-0776.1 (2014).Article 
PubMed 

Google Scholar 
Diez, J. M. et al. Will extreme climatic events facilitate biological invasions?. Front. Ecol. Environ. 10, 249–257. https://doi.org/10.1890/110137 (2012).Article 

Google Scholar 
Davis, M. A., Grime, J. P. & Thompson, K. Fluctuating resources in plant communities: a general theory of invasibility. J. Ecol. 88, 528–534. https://doi.org/10.1046/j.1365-2745.2000.00473.x (2000).Article 

Google Scholar 
Li, W. & Stevens, M. H. H. Fluctuating resource availability increases invasibility in microbial microcosms. Oikos 121, 435–441. https://doi.org/10.1111/j.1600-0706.2011.19762.x (2012).Article 

Google Scholar 
Koerner, S. E. et al. Invasibility of a mesic grassland depends on the time-scale of fluctuating resources. J. Ecol. 103, 1538–1546. https://doi.org/10.1111/1365-2745.12479 (2015).Article 

Google Scholar 
Hendrickson, J. R. & Lund, C. Plant community and target species affect responses to restoration strategies. Rangel. Ecol. Manag. 63, 435–442 (2010).Article 

Google Scholar 
Bennett, J., Smart, A. & Perkins, L. Using phenological niche separation to improve management in a Northern Glaciated Plains grassland. Restor. Ecol. 27, 745–749. https://doi.org/10.1111/rec.12932 (2019).Article 

Google Scholar 
Jordan, N. R., Larson, D. L. & Huerd, S. C. Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies. Biol. Invas. 10, 177–190. https://doi.org/10.1007/s10530-007-9121-1 (2008).Article 

Google Scholar 
Piper, C. L., Lamb, E. G. & Siciliano, S. D. Smooth brome changes gross soil nitrogen cycling processes during invasion of a rough fescue grassland. Plant Ecol. 216, 235–246. https://doi.org/10.1007/s11258-014-0431-y (2015).Article 

Google Scholar 
Stotz, G. C., Gianoli, E. & Cahill, J. F. Biotic homogenization within and across eight widely distributed grasslands following invasion by Bromus inermis. Ecology https://doi.org/10.1002/ecy.2717 (2019).Article 
PubMed 

Google Scholar 
Dillemuth, F. P., Rietschier, E. A. & Cronin, J. T. Patch dynamics of a native grass in relation to the spread of invasive smooth brome (Bromus inermis). Biol. Invas. 11, 1381–1391. https://doi.org/10.1007/s10530-008-9346-7 (2009).Article 

Google Scholar 
Trammell, M. A. & Butler, J. L. Effects of exotic plants on native ungulate use of habitat. J. Wildlife Manag. 59, 808–816. https://doi.org/10.2307/3801961 (1995).Article 

Google Scholar 
Gibson, D. J. Grasses and Grassland Ecology (Oxford Univ. Press, 2009).
Google Scholar 
Knapp, A. K. & Smith, M. D. Variation among biomes in temporal dynamics of aboveground primary production. Science 291, 481–484. https://doi.org/10.1126/science.291.5503.481 (2001).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Easterling, D. R. et al. Precipitation change in the United States. pp. 207–230 (Washington, D.C. USA, 2017).Gutschick, V. P. & BassiriRad, H. Extreme events as shaping physiology, ecology, and evolution of plants: toward a unified definition and evaluation of their consequences. New Phytol. 160, 21–42. https://doi.org/10.1046/j.1469-8137.2003.00866.x (2003).Article 
PubMed 

Google Scholar 
Briske, D. D. in Grazing management: An ecological perspective (eds R.K. Heitschmidt & J.W. Stuth) pp. 85–108 (Timber Press, Inc., 1991).Liu, F., Liu, J. & Dong, M. Ecological consequences of clonal integration in plants. Front. Plant Sci. 217, 277–287 (2016).
Google Scholar 
Hoover, D. L., Knapp, A. K. & Smith, M. D. Resistance and resilience of a grassland ecosystem to climate extremes. Ecology 95, 2646–2656. https://doi.org/10.1890/13-2186.1 (2014).Article 

Google Scholar 
VanderWeide, B. L., Hartnett, D. C. & Carter, D. L. Belowground bud banks of tallgrass prairie are insensitive to multi-year, growing-season drought. Ecosphere. https://doi.org/10.1890/Es14-00058.1 (2014).Article 

Google Scholar 
VanderWeide, B. L. & Hartnett, D. C. Belowground bud bank response to grazing under severe, short-term drought. Oecologia 178, 795–806. https://doi.org/10.1007/s00442-015-3249-y (2015).ADS 
Article 
PubMed 

Google Scholar 
Ott, J. P., Butler, J. L., Rong, Y. P. & Xu, L. Greater bud outgrowth of Bromus inermis than Pascopyrum smithii under multiple environmental conditions. J. Plant Ecol. 10, 518–527. https://doi.org/10.1093/jpe/rtw045 (2017).Article 

Google Scholar 
Oesterheld, M., Loreti, J., Semmartin, M. & Sala, O. E. Inter-annual variation in primary production of a semi-arid grassland related to previous-year production. J. Veg. Sci. 12, 137–142. https://doi.org/10.1111/j.1654-1103.2001.tb02624.x (2001).Article 

Google Scholar 
Ott, J. P. & Hartnett, D. C. Bud bank dynamics and clonal growth strategy in the rhizomatous grass, Pascopyrum smithii. Plant Ecol. 216, 395–405. https://doi.org/10.1007/s11258-014-0444-6 (2015).Article 

Google Scholar 
Carlsson, B. A. & Callaghan, T. V. Programmed tiller differentiation, intraclonal density regulation and nutrient dynamics in Carex bigelowii. Oikos 58, 219–230. https://doi.org/10.2307/3545429 (1990).Article 

Google Scholar 
Ye, X. H., Yu, F. H. & Dong, M. A trade-off between guerrilla and phalanx growth forms in Leymus secalinus under different nutrient supplies. Ann. Bot. Lond. 98, 187–191. https://doi.org/10.1093/aob/mcl086 (2006).Article 

Google Scholar 
Dibbern, J. C. Vegetative responses of Bromus inermis to certain variations in environment. Bot. Gazette 109, 44–58 (1947).Article 

Google Scholar 
Dong, X., Patton, J., Wang, G., Nyren, P. & Peterson, P. Effect of drought on biomass allocation in two invasive and two native grass species dominating the mixed-grass prairie. Grass Forage Sci. 69, 160–166. https://doi.org/10.1111/gfs.12020 (2014).Article 

Google Scholar 
Saeidnia, F., Majidi, M. M., Mirlohi, A. & Soltan, S. Physiological and tolerance indices useful for drought tolerance selection in smooth bromegrass. Crop Sci. 57, 282–289. https://doi.org/10.2135/cropsci2016.07.0636 (2017).CAS 
Article 

Google Scholar 
Vinton, M. A. & Hartnett, D. C. Effects of bison grazing on Andropogon gerardii and Panicum virgatum in burned and unbruned tallgrass prairie. Oecologia 90, 374–382. https://doi.org/10.1007/bf00317694 (1992).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Eneboe, E. J., Sowell, B. F., Heitschmidt, R. K., Karl, M. G. & Haferkamp, M. R. Drought and grazing: IV. Blue grama and western wheatgrass. J. Range Manag. 55, 197–203. https://doi.org/10.2307/4003357 (2002).Article 

Google Scholar 
Broadbent, T. S., Bork, E. W. & Willms, W. D. Divergent effects of defoliation intensity and frequency on tiller growth and production dynamics of Pascopyrum smithii and Hesperostipa comata. Grass Forage Sci. 73, 532–543. https://doi.org/10.1111/gfs.12318 (2018).Article 

Google Scholar 
Donkor, N. T., Bork, E. W. & Hudson, R. J. Bromus-Poa response to defoliation intensity and frequency under three soil moisture levels. Can. J. Plant Sci. 82, 365–370. https://doi.org/10.4141/p01-040 (2002).Article 

Google Scholar 
Reynolds, J. H. & Smith, D. Trend of carbohydrate reserves in alfalfa, smooth bromegrass, and timothy grown under various cutting schedules. Crop Sci. 2, 333–336 (1962).CAS 
Article 

Google Scholar 
Lamp, H. F. Reproductive activity in Bromus inermis in relation to phases of tiller development. Bot. Gazette 113, 413–438 (1952).Article 

Google Scholar 
Paulsen, G. M. & Smith, D. Organic reserves, axillary bud activity, and herbage yields of smooth bromegrass as influenced by time of cutting, nitrogen fertilization, and shading. Crop Sci. 9, 529–534 (1969).Article 

Google Scholar 
Ott, J. P. & Hartnett, D. C. Contrasting bud bank dynamics of two co-occurring grasses in tallgrass prairie: implications for grassland dynamics. Plant Ecol. 213, 1437–1448. https://doi.org/10.1007/s11258-012-0102-9 (2012).Article 

Google Scholar 
Busso, C. A., Mueller, R. J. & Richards, J. H. Effects of drought and defoliation on bud viability in 2 caespitose grasses. Ann. Bot. Lond. 63, 477–485. https://doi.org/10.1093/oxfordjournals.aob.a087768 (1989).Article 

Google Scholar 
Tuomi, J., Nilsson, P. & Astrom, M. Plant compensatory responses-bud dormancy as an adaptation to herbivory. Ecology 75, 1429–1436. https://doi.org/10.2307/1937466 (1994).Article 

Google Scholar 
US Department of Agriculture. The PLANTS Database, (2006).Gong, K. et al. Analysis on the distribution, breeding and utilization of Bromus inermis germplasm resource in China. Heilongjiang Anim. Sci. Vet. Med. 21, 33–36 (2019).
Google Scholar 
Coupland, R. T. & Johnson, R. E. Rooting characteristics of native grassland species in Saskatchewan. J. Ecol. 53, 475–507 (1965).Article 

Google Scholar 
Gist, G. R. & Smith, R. M. Root development of several common forage grasses to a depth of eighteen inches. Agron. J. 1036–1042 (1948).Okamoto, H., Ishii, K. & An, P. Effects of soil moisture deficit and subsequent watering on the growth of four temperate grasses. Grassl. Sci. 57, 192–197. https://doi.org/10.1111/j.1744-697X.2011.00232.x (2011).Article 

Google Scholar 
Morrow, L. A. & Power, J. F. Effect of soil temperature on development of perennial forage grasses. Agron. J. 71, 7–10 (1979).Article 

Google Scholar 
Duell, E. B., Wilson, G. W. T. & Hickman, K. R. Above- and below-ground responses of native and invasive prairie grasses to future climate scenarios. Botany 94, 471–479. https://doi.org/10.1139/cjb-2015-0238 (2016).Article 

Google Scholar 
Duell, E. B., Londe, D. W., Hickman, K. R., Greer, M. J. & Wilson, G. W. T. Superior performance of invasive grasses over native counterparts will remain problematic under warmer and drier conditions. Plant Ecol. 222, 993–1006 (2021).Article 

Google Scholar 
Cully, A. C., Cully, J. F. & Hiebert, R. D. Invasion of exotic plant species in tallgrass prairie fragments. Conser. Biol. 17, 990–998. https://doi.org/10.1046/j.1523-1739.2003.02107.x (2003).Article 

Google Scholar 
DeKeyser, E. S., Meehan, M., Clambey, G. & Krabbenhoft, K. Cool season invasive grasses in northern great plains natural areas. Nat. Areas J. 33, 81–90. https://doi.org/10.3375/043.033.0110 (2013).Article 

Google Scholar 
Grant, T. A., Shaffer, T. L. & Flanders, B. Resiliency of native prairies to invasion by kentucky bluegrass, smooth brome, and woody vegetation. Rangeland Ecol. Manag. 73, 321–328. https://doi.org/10.1016/j.rama.2019.10.013 (2020).Article 

Google Scholar 
Otfinowski, R., Kenkel, N. C. & Catling, P. M. The biology of Canadian weeds. 134. Bromus inermis Leyss. Can. J. Plant Sci. 87, 183–198. https://doi.org/10.4141/p06-071 (2007).Article 

Google Scholar 
Moore, K. J. et al. Describing and quantifying growth stages of perennial forage grasses. Agron. J. 83, 1073–1077 (1991).Article 

Google Scholar 
SAS Institute. SAS 9.4. (SAS Institute Inc, 2017). More

Phillips, R. et al. The conservation status and priorities for albatrosses and large petrels. Biol. Conserv. 201, 169–183. https://doi.org/10.1016/j.biocon.2016.06.017 (2016).Article 

Google Scholar 
Dias, M. et al. Threats to seabirds: A global assessment. Biol. Conserv. 237, 525–537. https://doi.org/10.1016/j.biocon.2019.06.033 (2019).Article 

Google Scholar 
IUCN. The IUCN Red List of Threatened Species. Version 2021-1, www.iucnredlist.org, ISSN 2307-8235. (International Union for the Conservation of Nature, 2021).Brothers, N., Cooper, J. & Løkkeborg, S. The Incidental Catch of Seabirds by Longline Fisheries: Worldwide Review and Technical Guidelines for Mitigation. FAO Fisheries Circular No. 937. (Food and Agriculture Organization of the United Nations, 1999)Gilman, E., Brothers, N. & Kobayashi, D. Principles and approaches to abate seabird bycatch in longline fisheries. Fish Fish. 6, 35–49. https://doi.org/10.1111/j.1467-2679.2005.00175.x (2005).Article 

Google Scholar 
Løkkeborg, S. Best practices to mitigate seabird by—catch in longline, trawl and gillnet fisheries—efficiency and practical applicability. Mar. Ecol. Prog. Ser. 435, 285–303. https://doi.org/10.3354/meps09227 (2011).ADS 
Article 

Google Scholar 
Gilman, E., Chaloupka, M., Peschon, J. & Ellgen, S. Risk factors for seabird bycatch in a pelagic longline tuna fishery. PLoS One 11, e0155477 (2016).Article 

Google Scholar 
Gilman, E., Kobayashi, D. & Chaloupka, M. Reducing seabird bycatch in the Hawaii longline tuna fishery. Endanger. Species Res. 5, 309–323. https://doi.org/10.3354/esr00133 (2008).Article 

Google Scholar 
WPRFMC. Annual Stock Assessment and Fishery Evaluation Report for U.S. Pacific Island Pelagic Fisheries Ecosystem Plan 2019. (Western Pacific Regional Fishery Management Council, Honolulu, 2020).Wren, J., Shaffer, S. & Polovina, J. Variations in black-footed albatross sightings in a North Pacific transition area due to changes in fleet dynamics and oceanography 2006–2017. Deep. Sea. Res. Part II Top. Stud. Oceanogr. 169–170, 104605. https://doi.org/10.1016/j.dsr2.2019.06.013 (2019).Article 

Google Scholar 
NMFS. Seabird Interactions and Mitigation Efforts in Hawaii Longline Fisheries. 2019 Annual Report. (Pacific Islands Regional Office, National Marine Fisheries Service, 2021).Hall, M., Gilman, E., Minami, H., Mituhasi, T. & Carruthers, E. Mitigating bycatch in tuna fisheries. Rev. Fish. Biol. Fisher. 27, 881–908. https://doi.org/10.1007/s11160-017-9478-x (2017).Article 

Google Scholar 
ACAP. ACAP Review and Best Practice Advice for Reducing the Impact of Pelagic Longline Fisheries on Seabirds. (Agreement on the Conservation of Albatrosses and Petrels, 2019).NMFS. Fisheries off west coast states and in the western Pacific; pelagic fisheries; additional measures to reduce the incidental catch of seabirds in the Hawaii pelagic longline fishery. US National Marine Fisheries Service. Fed. Regist. 70, 75075–77508 (2005).
Google Scholar 
Robertson, G., Candy, S. & Hall, S. New branch line weighting regimes to reduce the risk of seabird mortality in pelagic longline fisheries without affecting fish catch. Aquat. Conserv. 23, 885–900. https://doi.org/10.1002/aqc.2346 (2013).Article 

Google Scholar 
Melvin, E., Guy, T. & Read, L. Reducing seabird bycatch in the South African tuna fishery using bird-scaring lines, branch line weighting and nighttime setting of hooks. Fish. Res. 147, 72–82 (2013).Article 

Google Scholar 
Melvin, E., Guy, T. & Read, L. Best practice seabird bycatch mitigation for pelagic longline fisheries targeting tuna and related species. Fish. Res. 149, 5–18 (2014).Article 

Google Scholar 
Santos, R. et al. Improved line weighting reduces seabird bycatch without affecting fish catch in the Brazilian pelagic longline fishery. Aquat. Conserv. 29, 442–449. https://doi.org/10.1002/aqc.3002 (2019).Article 

Google Scholar 
Gilman, E., Chaloupka, M., Wiedoff, B. & Willson, J. Mitigating seabird bycatch during hauling by pelagic longline vessels. PLoS One 9, e84499. https://doi.org/10.1371/journal.pone.0084499 (2014).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Brothers, N. Incidence of Live Bird Haul Capture in Pelagic Longline Fisheries. Examination and Comparison of Live Bird Haul Captures in Fisheries Other Than the Hawaii Shallow Set Fishery Agreement on the Conservation of Albatrosses and Petrels. SBWG7 Doc 18. (Agreement on the Conservation of Albatrosses and Petrels, 2016).Jiminez, S., Domingo, A., Forselledo, R., Sullivan, B. & Yates, O. Mitigating bycatch of threatened seabirds: The effectiveness of branch line weighting in pelagic longline fisheries. Anim. Conserv. 22, 376–385. https://doi.org/10.1111/acv.12472 (2019).Article 

Google Scholar 
Bentley, L., Kato, A., Ropert-Coudert, Y., Manica, A. & Phillips, R. Diving behaviour of albatrosses: Implications for foraging ecology and bycatch susceptibility. Mar. Biol. 168, 36. https://doi.org/10.1007/s00227-021-03841-y (2021).Article 

Google Scholar 
Prince, P., Huin, N. & Weimerskirch, H. Diving depths of albatrosses. Antarct. Sci. 6, 353–354. https://doi.org/10.1017/S0954102094000532 (1994).ADS 
Article 

Google Scholar 
Kazama, K., Harada, T., Deguchi, T., Suzuki, H. & Watanuki, Y. Foraging behavior of black-footed albatross Phoebastria nigripes rearing chicks on the Ogasawara Islands. Ornithol. Sci. 18, 27–37 (2019).Article 

Google Scholar 
Jiminez, S., Domingo, A., Abreu, M. & Brazeiro, A. Bycatch susceptibility in pelagic longline fisheries: Are albatrosses affected by the diving behaviour of medium-sized petrels?. Aquat. Conserv. 22, 436–445. https://doi.org/10.1002/aqc.2242 (2012).Article 

Google Scholar 
Barrington, J., Robertson, G. & Candy, S. Categorising Branchline Weighting for Pelagic Longline Fishing According to Sink Rate. ACAP-SBWG7-Doc7. (Agreement on the Conservation of Albatrosses and Petrels, 2016).NOAA. Endangered and threatened wildlife and plants: Listing the oceanic whitetip shark as threatened under the Endangered Species Act. Fed. Regist. 83, 4153–4165 (2018).
Google Scholar 
WPRFMC. Council Adopts Oceanic Whitetip Shark Protections for Hawaii and American Samoa Longline Fisheries. (Western Pacific Regional Fishery Management Council, 2021).Pierre, J., Goad, D. & Abraham, E. Novel Approaches to Line-Weighting in New Zealand’s Inshore Surface-Longline Fishery. (Dragonfly Data Science, 2015).Rawlinson, N., et al. The Relative Safety of Weighted Branchlines During Simulated Fly-backs (Cut-offs and Tear-outs). (AMC Research, 2018).Gilman, E., Beverly, S., Musyl, M. & Chaloupka, M. Commercial viability of alternative designs placing pelagic longline branchline weights at the hook to reduce seabird bycatch. Endanger. Species Res. 43, 223–233 (2020).Article 

Google Scholar 
Fenaughty, J. & Smith, N. A Simple New Method for Monitoring Longline Sink Rate to Selected Depths. Document WG-FSA-01/46. (Commission for the Conservation of Antarctic Marine Living Resources, 2001).Wienecke, B. & Robertson, G. Validation of sink rates of longlines measured using two different methods. CCAMLR Sci. 11, 179–187 (2004).
Google Scholar 
Melvin, E. & Wainstein, M. Seabird Avoidance Measures for Small Alaskan Longline Vessels (University of Washington, 2006).
Google Scholar 
CCAMLR. Longline Weighting for Seabird Conservation. Conservation Measure 24-02. (Commission for the Conservation of Antarctic Marine Living Resources, 2014).Robertson, G., Candy, S., Wienecke, B. & Lawton, K. Experimental determinations of factors affecting the sink rates of baited hooks to minimize seabird mortality in pelagic longline fisheries. Aquat. Conserv. 20, 632–643. https://doi.org/10.1002/aqc.1140 (2010).Article 

Google Scholar 
Wondershare Technology. Wondershare Filmora X. Version 10.2.0.32. (Wondershare Technology Co., 2021).Gabry, J., Simpson, D., Vehtari, A., Betancourt, M. & Gelman, A. Visualization in Bayesian workflow. J. R. Stat. Soc. Ser. A 182, 1–14. https://doi.org/10.1111/rssa.12378 (2019).MathSciNet 
Article 

Google Scholar 
Gelman, A., et al. Bayesian Workflow. arXiv:2011.01808v1 (2020).Gelman, A, & Hill, J. Data Analysis Using Regression and Multilevel/Hierarchical Models. (Cambridge University Press, 2007).Carpenter, B. et al. Stan: A probabilistic programming language. J. Stat. Softw. 76, 1–32. https://doi.org/10.18637/jss.v076.i01 (2017).Article 

Google Scholar 
Bürkner, P. brms: An R Package for Bayesian multilevel models using Stan. J. Stat. Softw. 81, 1–28. https://doi.org/10.18637/jss.v080.i01 (2017).Article 

Google Scholar 
Gilman, E., Chaloupka, M. & Musyl, M. Effects of pelagic longline hook size on species- and size-selectivity and survival. Rev. Fish. Biol. Fish. 28, 417–433. https://doi.org/10.1007/s11160-017-9509-7 (2018).Article 

Google Scholar 
Signorell, A., et al. DescTools: Tools for Descriptive Statistics. R package version 0.99.18. (R Core Team, 2016).Wackerly, D., Mendenhall, W. & Scheaffer, R. Mathematical Statistics with Applications. 3rd ed. (Duxbury Press, 1986).van Houwelingen, H., Arends, L. & Stijnen, T. Advanced methods in meta-analysis: Multivariate approach and meta-regression. Stat. Med. 21, 589–624. https://doi.org/10.1002/sim.1040 (2002).Article 
PubMed 

Google Scholar 
Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Softw. 36, 1–48. https://doi.org/10.18637/jss.v036.i03 (2010).Article 

Google Scholar 
Günhan, B., Röver, C. & Friede, T. Random-effects meta-analysis of few studies involving rare events. Res. Synth. Methods 11, 74–90. https://doi.org/10.1002/jrsm.1370 (2020).Article 
PubMed 

Google Scholar 
Lemoine, N. Moving beyond noninformative priors: Why and how to choose weakly informative priors in Bayesian analyses. Oikos 128, 912–928. https://doi.org/10.1111/oik.05985 (2019).Article 

Google Scholar 
Schild, A. & Voracek, M. Finding your way out of the forest without a trail of bread crumbs: Development and evaluation of two novel displays of forest plots. Res. Synth. Methods 6, 74–86. https://doi.org/10.1002/jrsm.1125 (2015).Article 
PubMed 

Google Scholar 
van der Bles, A. et al. Communicating uncertainty about facts, numbers and science. R. Soc. Open Sci. 6, 181870. https://doi.org/10.1098/rsos.181870 (2019).ADS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Wickham, H. ggplot2: Elegant Graphics for Data Analysis. 2nd ed. (Springer, 2016).Zeileis, A. et al. colorspace: A toolbox for manipulating and assessing colors and palettes. J. Stat. Softw. 96, 1–4. https://doi.org/10.18637/jss.v096.i01 (2020).Article 

Google Scholar 
Vovk, V. & Wang, R. Combining p-values via averaging. Biometrika 107, 791–808. https://doi.org/10.1093/biomet/asaa027 (2020).MathSciNet 
Article 
MATH 

Google Scholar 
Gilman, E., Brothers, N. & Kobayashi, D. Comparison of the efficacy of three seabird bycatch avoidance methods in Hawaii pelagic longline fisheries. Fish. Sci. 73, 208–210 (2007).CAS 
Article 

Google Scholar 
Star-Oddi. DST centi-TD Miniature Temperature and Depth Data Logger. (Star-Oddi, 2021).Frankish, C., Manica, A., Navarro, J. & Phillips, R. Movements and diving behaviour of white-chinned petrels: Diurnal variation and implications for bycatch mitigation. Aquat. Conserv. 31, 1715–1729. https://doi.org/10.1002/aqc.3573 (2021).Article 

Google Scholar 
Cooke, S. & Suski, C. Are circular hooks and effective tool for conserving marine and freshwater recreational catch-and-release fisheries?. Aquat. Conserv. 14, 299–326. https://doi.org/10.1002/aqc.614 (2004).Article 

Google Scholar 
Ward, P., Lawrence, E., Darbyshire, R. & Hindmarsh, S. Large-scale experiment shows that nylon leaders reduce shark bycatch and benefit pelagic longline fishers. Fish. Res. 90, 100–108. https://doi.org/10.1016/j.fishres.2007.09.034 (2008).Article 

Google Scholar 
McCormack, E. & Rawlinson, N. The Relative Safety of the Agreement on the Conservation of Albatrosses and Petrels (ACAP) Recommended Minimum Specifications for the Weighting of Branchlines during Simulated Fly-backs. ACAP-SBWG7-Doc8. (Agreement on the Conservation of Albatrosses and Petrels, 2016).Goad, D., Debski, I. & Potts, J. Hookpod-mini: A smaller potential solution to mitigate seabird bycatch in pelagic longline fisheries. Endanger. Species Res. 39, 1–8. https://doi.org/10.3354/esr00953 (2019).Article 

Google Scholar 
WHO. Global Health Risks: Mortality and Burden of Disease Attributable to Selected Major Risks. (World Health Organization, 2009).Grade, T. et al. Lead poisoning from ingestion of fishing gear: A review. Ambio 48, 1023–1038. https://doi.org/10.1007/s13280-019-01179-w (2019).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Gilman, E. et al. Highest risk abandoned, lost and discarded fishing gear. Sci. Rep. 11, 7195. https://doi.org/10.1038/s41598-021-86123-3 (2021).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Brothers, N. In Pursuit of Procella—A Heavy Hook for Pelagic Longlines to Reduce Procellariiforme Bycatch. SBWG 10 Inf 09. (Agreement on the Conservation of Albatrosses and Petrels, 2021).Bluhm, R. From hierarchy to network: A richer view of evidence for evidence-based medicine. Perspect. Biol. Med. 48, 535–547. https://doi.org/10.1353/pbm.2005.0082 (2005).Article 
PubMed 

Google Scholar 
Stegenga, J. Down with the hierarchies. Topoi 33, 313–322. https://doi.org/10.1007/s11245-013-9189-4 (2014).Article 

Google Scholar 
Marchionni, C. & Reijula, S. What is mechanistic evidence, and why do we need it for evidence-based policy?. Stud. Hist. Philos. Sci. A 73, 54–63. https://doi.org/10.1016/j.shpsa.2018.003 (2019).Article 

Google Scholar 
Beverly, S., Chapman, L. & Sokimi, W. Horizontal Longline Fishing Methods and Techniques. Manual for Fishermen. (Secretariat of the Pacific Community, 2003).Guilford, T., Padget, O., Maurice, L. & Catry, P. Unexpectedly deep diving in an albatross. Curr. Biol. 32, R26–R28. https://doi.org/10.1016/j.cub.2021.11.036 (2022).CAS 
Article 
PubMed 

Google Scholar  More

Ebner, J., Babbitt, C., Winer, M., Hilton, B. & Williamson, A. Life cycle greenhouse gas (GHG) impacts of a novel process for converting food waste to ethanol and co-products. Appl. Energy 130, 86–93 (2014).CAS 

Google Scholar 
Tonini, D., Albizzati, P. F. & Astrup, T. F. Environmental impacts of food waste: Learnings and challenges from a case study on UK. Waste Manag. 76, 744 (2018).PubMed 

Google Scholar 
Sze, E., Yau, Y. H. & Wu, K. C. Application of anaerobic bacterial ammonification pretreatment to microalgal food waste leachate cultivation and biofuel production. Mar. Pollut. Bull. 153, 111007 (2020).PubMed 

Google Scholar 
Winkel, T. D., Wahlen, S. & Jensen, T. in Nordic Conference on Consumer Research.Wang, P. et al. Effects of graphite, graphene, and graphene oxide on the anaerobic co-digestion of sewage sludge and food waste: attention to methane production and the fate of antibiotic resistance genes. Bioresour. Technol. 339, 125585 (2021).CAS 
PubMed 

Google Scholar 
Gianico, A., Gallipoli, A., Pagliaccia, P. & Braguglia, C. M. Anaerobic bioconversion of food waste into energy: A critical review (2013).Smetana, S., Ites, S., Parniakov, O., Aganovic, K. & Heinz, V. in 71st Annual Meeting of the European Federation of Animal Science.Ojha, S., Buler, S. & Schlüter, O. Food waste valorisation and circular economy concepts in insect production and processing. Waste Manag. 118, 600–609 (2020).CAS 
PubMed 

Google Scholar 
Scala, A. et al. Rearing substrate impacts growth and macronutrient composition of Hermetia illucens (L.) (Diptera: Stratiomyidae) larvae produced at an industrial scale. Sci. Rep. 10, 1–8 (2020).ADS 

Google Scholar 
McDonald, C., Campbell, K. A., Benson, C., Davis, M. J. & Frost, C. J. Workforce development and multiagency collaborations: a presentation of two case studies in child welfare. Sustainability 13, 10190 (2021).
Google Scholar 
Kim, C.-H. et al. Use of black soldier fly larvae for food waste treatment and energy production in asian countries: a review. Processes 9, 161 (2021).CAS 

Google Scholar 
Julita, U., Fitri, L., Putra, R. & Permana, A. Mating success and reproductive behavior of black soldier fly Hermetia illucens L. (diptera, stra-tiomyidae) in tropics. J. Ento-mol. 17, 117–127 (2020).CAS 

Google Scholar 
Rehman, K. U. et al. Conversion of mixtures of dairy manure and soybean curd residue by black soldier fly larvae (Hermetia illucens L.). J. Clean. Prod. 154, 366–373 (2017).CAS 

Google Scholar 
Li, Q., Zheng, L., Hao, C., Garza, E. & Zhou, S. From organic waste to biodiesel: Black soldier fly, Hermetia illucens, makes it feasible. Fuel 90, 1545–1548 (2011).CAS 

Google Scholar 
Köhler, R., Kariuki, L., Lambert, C. & Biesalski, H. K. Protein, amino acid and mineral composition of some edible insects from Thailand. J. Asia Pac. Entomol. 22, 372–378 (2019).
Google Scholar 
Belghit, I. et al. Black soldier fly larvae meal can replace fish meal in diets of sea-water phase Atlantic salmon (Salmo salar). Aquaculture (2018).Moretta, A. et al. Antimicrobial peptides: A new hope in biomedical and pharmaceutical fields. Front. Cell. Infect. Microbiol. 11, 453 (2021).
Google Scholar 
Manniello, M. et al. Insect antimicrobial peptides: potential weapons to counteract the antibiotic resistance. Cell. Mol. Life Sci. 89, 1–24 (2021).
Google Scholar 
Henriques, B. S., Garcia, E. S., Azambuja, P. & Genta, F. A. Determination of chitin content in insects: an alternate method based on calcofluor staining. Front. Physiol. 11, ARTN 11710.3389/fphys.2020.00117 (2020).Hbl, M., Mráz, P., Ipo, J., Hotiková, I. & Kopec, T. Polyphenols as food supplement improved food consumption and longevity of honey bees (Apis mellifera) intoxicated by pesticide thiacloprid. Insects 12 (2021).Li, H., Dai, C., Zhu, Y. & Hu, Y. Larvae crowding increases development rate, improves disease resistance, and induces expression of antioxidant enzymes and heat shock proteins in Mythimna separata (Lepidoptera: Noetuidae). J. Econ. Entomol. 4 (2021).Hao, et al. Effects of enzymatic hydrolysis assisted by high hydrostatic pressure processing on the hydrolysis and allergenicity of proteins from ginkgo seeds. Food Bioprocess Technol. 9, 839–848 (2016).
Google Scholar 
Nadeem, M., Mumtaz, M. W., Danish, M., Rashid, U. & Raza, S. A. Calotropis procera: UHPLC-QTOF-MS/MS based profiling of bioactives, antioxidant and anti-diabetic potential of leaf extracts and an insight into molecular docking. J. Food Meas. Charact. 13, 3206–3220 (2019).
Google Scholar 
Altomare, A. A., Baron, G., Aldini, G., Carini, M. & D’Amato, A. Silkworm pupae as source of high-value edible proteins and of bioactive peptides. Food Sci. Nutr. 8, 2652–2661 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Zielińska, E., Baraniak, B. & Karaś, M. Identification of antioxidant and anti-inflammatory peptides obtained by simulated gastrointestinal digestion of three edible insects species (Gryllodes sigillatus, Tenebrio molitor, Schistocerca gragaria). Int. J. Food Sci. Technol. 53, 2542–2551 (2018).
Google Scholar 
Sousa, P., Borges, S. & Pintado, M. Enzymatic hydrolysis of insect Alphitobius diaperinus towards the development of bioactive peptide hydrolysates. Food Funct. 11, 3539–3548 (2020).CAS 
PubMed 

Google Scholar 
Jakubczyk, A., Karaś, M., Rybczyńska-Tkaczyk, K., Zielińska, E. & Zieliński, D. Current trends of bioactive peptides—New sources and therapeutic effect. Foods 9, 846 (2020).CAS 
PubMed Central 

Google Scholar 
Li, K., Li, X.-M., Ji, N.-Y. & Wang, B.-G. Natural bromophenols from the marine red alga Polysiphonia urceolata (Rhodomelaceae): structural elucidation and DPPH radical-scavenging activity. Bioorg. Med. Chem. 15, 6627–6631 (2007).CAS 
PubMed 

Google Scholar 
He, R., Girgih, A. T., Malomo, S. A., Ju, X. R. & Aluko, R. E. Antioxidant activities of enzymatic rapeseed protein hydrolysates and the membrane ultrafiltration fractions. J. Funct. Foods 5, 219–227. https://doi.org/10.1016/j.jff.2012.10.008 (2013).CAS 
Article 

Google Scholar 
Cui, Q., Sun, Y. X., Cheng, J. J. & Guo, M. R. Effect of two-step enzymatic hydrolysis on the antioxidant properties and proteomics of hydrolysates of milk protein concentrate. Food Chem. 366, 10. https://doi.org/10.1016/j.foodchem.2021.130711 (2022).CAS 
Article 

Google Scholar 
Liu, Y., Wan, S., Liu, J., Zou, Y. & Liao, S. Antioxidant activity and stability study of peptides from enzymatically hydrolyzed male silkmoth. J. Food Process. Preserv. 41 (2017).Carrasco-Castilla, J. et al. Antioxidant and metal chelating activities of peptide fractions from phaseolin and bean protein hydrolysates. Food Chem. 135, 1789–1795 (2012).CAS 
PubMed 

Google Scholar 
Phongthai, S., D’Amico, S., Schoenlechner, R., Homthawornchoo, W. & Rawdkuen, S. Fractionation and antioxidant properties of rice bran protein hydrolysates stimulated by in vitro gastrointestinal digestion. Food Chem. 240, 156 (2018).CAS 
PubMed 

Google Scholar 
Lee, S. J. et al. Antioxidant activity of a novel synthetic hexa-peptide derived from an enzymatic hydrolysate of duck skin by-products. Food Chem. Toxicol. 62, 276–280 (2013).CAS 
PubMed 

Google Scholar 
Collin, F. Chemical basis of reactive oxygen species reactivity and involvement in neurodegenerative diseases. Int. J. Mol. Sci. 20, 2407 (2019).PubMed Central 

Google Scholar 
Xiang, Q., Yu, J. & Wong, P. K. Quantitative characterization of hydroxyl radicals produced by various photocatalysts. J. Colloid Interface Sci. 357, 163–167 (2011).ADS 
CAS 
PubMed 

Google Scholar 
Wang, Y. et al. Optimizing oxygen functional groups in graphene quantum dots for improved antioxidant mechanism. Phys. Chem. Chem. Phys. 21, 1336–1343 (2019).CAS 
PubMed 

Google Scholar 
Arise, A. K. et al. Antioxidant activities of bambara groundnut (Vigna subterranea) protein hydrolysates and their membrane ultrafiltration fractions. Food Funct. 7, 2431–2437. https://doi.org/10.1039/c6fo00057f (2016).CAS 
Article 
PubMed 

Google Scholar 
Ren, J. et al. Purification and identification of antioxidant peptides from grass carp muscle hydrolysates by consecutive chromatography and electrospray ionization-mass spectrometry. Food Chem. 108, 727–736. https://doi.org/10.1016/j.foodchem.2007.11.010 (2008).CAS 
Article 
PubMed 

Google Scholar 
Zhang, C., Wei, X., Omenn, G. S. & Zhang, Y. Structure and protein interaction-based gene ontology annotations reveal likely functions of uncharacterized proteins on human chromosome 17. 17 (2018).Long, C. N. et al. High-level production of Monascus pigments in Monascus ruber CICC41233 through ATP-citrate lyase overexpression. Biochem. Eng. J. 146, 160–169. https://doi.org/10.1016/j.bej.2019.03.007 (2019).CAS 
Article 

Google Scholar 
Brito Querido, J. et al. The cryo-EM structure of a novel 40S kinetoplastid-specific ribosomal protein. Structure 25, 1785–1794 e1783. https://doi.org/10.1016/j.str.2017.09.014 (2017).Hamey, J. J. & Wilkins, M. R. Methylation of elongation factor 1A: Where, who, and why?. Trends Biochem. Sci. 43, 211–223. https://doi.org/10.1016/j.tibs.2018.01.004 (2018).CAS 
Article 
PubMed 

Google Scholar 
Kuo, C. P. et al. Analysis of the immune response of human dendritic cells to Mycobacterium tuberculosis by quantitative proteomics. Proteome Sci. 14, 1–11 (2016).
Google Scholar 
Zhu, J. et al. Expression and RNA interference of ribosomal protein L5 gene in Nilaparvata lugens (Hemiptera: Delphacidae). J. Insect. Sci. 3 (2017).Teng, T., Mercer, C. A., Hexley, P., Thomas, G. & Fumagalli, S. Loss of tumor suppressor RPL5/RPL11 does not induce cell cycle arrest but impedes proliferation due to reduced ribosome content and translation capacity. Mol. Cell. Biol. 33, 4660–4671. https://doi.org/10.1128/mcb.01174-13 (2013).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Rittschof, C. C. & Schirmeier, S. Insect models of central nervous system energy metabolism and its links to behavior. Glia (2017).Alar, A. F., Akr, B. & Gülseren, I. LC-Q-TOF/MS based identification and in silico verification of ACE-inhibitory peptides in Giresun (Turkey) hazelnut cakes. Eur. Food Res. Technol. (2021).Shang, W. H. et al. In silico assessment and structural characterization of antioxidant peptides from major yolk protein of sea urchin Strongylocentrotus nudus. Food Funct. 9, 6435–6443 (2018).CAS 
PubMed 

Google Scholar 
Ajibola, C. F., Fashakin, J. B., Fagbemi, T. N. & Aluko, R. E. Effect of peptide size on antioxidant properties of African yam bean seed (Sphenostylis stenocarpa) protein hydrolysate fractions. Int. J. Mol. Sci. 12, 6685–6702 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Liu, H. et al. Enhancing the antioxidative effects of foods containing rutin and α-amino acids via the Maillard reaction: A model study focusing on rutin-lysine system. J. Food Biochem. 44, e13086 (2020).PubMed 

Google Scholar 
Tsopmo, A. et al. Tryptophan released from mother’s milk has antioxidant properties. Pediatr. Res. 66, 614–618 (2009).CAS 
PubMed 

Google Scholar 
Yu, Z. et al. Identification and molecular docking study of fish roe-derived peptides as potent BACE 1, AChE, and BChE inhibitors. Food Funct. 11 (2020).Li, C. et al. Preliminary study on a potential antibacterial peptide derived from histone H2A in hemocytes of scallop Chlamys farreri. Fish Shellf. Immunol. 22, 663–672 (2007).
Google Scholar 
Arockiaraj, J. et al. An unconventional antimicrobial protein histone from freshwater prawn Macrobrachium rosenbergii: analysis of immune properties. Fish Shellfish Immunol. 35, 1511–1522 (2013).CAS 
PubMed 

Google Scholar 
Ju, J. et al. Major components in Lilac and Litsea cubeba essential oils kill Penicillium roqueforti through mitochondrial apoptosis pathway. Ind. Crops Products 149, 112349 (2020).CAS 

Google Scholar 
Al-Dhafri, K., Chai, L. C. & Karsani, S. A. Purification and characterization of antimicrobial peptide fractions of Junipers seravschanica. Biocatal. Agric. Biotechnol. 28, 101554 (2020).
Google Scholar 
Ratnakomala, S., Ridwan, R., Lisdiyanti, P., Abinawanto, A. & Andi, U. Screening of actinomycetes producing an ATPase inhibitor of japanese encephalitis virus RNA helicase from soil and leaf litter samples. Microbiol. Indonesia 5, 15–20 (2011).
Google Scholar 
Zhao, X., Zhang, J. & Zhu, K. Y. Chito-protein matrices in arthropod exoskeletons and peritrophic matrices (2019).Pustylnikov, S., Sagar, D., Jain, P. & Khan, Z. K. Targeting the C-type lectins-mediated host-pathogen interactions with dextran. J. Pharm. Pharm. Sci. 17 (2014).Kiew, P. L. & Don, M. M. Jewel of the seabed: sea cucumbers as nutritional and drug candidates. Int. J. Food Sci. Nutr. 63, 616–636 (2012).CAS 
PubMed 

Google Scholar 
Liu, Z., Su, Y. & Zeng, M. Amino acid composition and functional properties of giant red sea cucumber (Parastichopus californicus) collagen hydrolysates. J. Ocean Univ. China 10, 80–84 (2011).ADS 
CAS 

Google Scholar 
Zaky, A. A., Liu, Y., Han, P., Chen, Z. & Jia, Y. Effect of pepsin–trypsin in vitro gastro-intestinal digestion on the antioxidant capacities of ultra-filtrated rice bran protein hydrolysates (molecular weight > 10 kDa; 3–10 kDa, and< 3 kDa). Int. J. Pept. Res. Ther. 1–7 (2019).Kim, S.-B., Yoon, N. Y., Shim, K.-B. & Lim, C.-W. Antioxidant and angiotensin I-converting enzyme inhibitory activities of northern shrimp (Pandalus borealis) by-products hydrolysate by enzymatic hydrolysis. Fish. Aquat. Sci. 19, 1–6 (2016). Google Scholar  Chung, Y. C., Chang, C. T., Chao, W. W., Lin, C. F. & Chou, S. T. Antioxidative activity and safety of the 50 ethanolic extract from red bean fermented by Bacillus subtilis IMR-NK1. J. Agric. Food Chem. 50, 2454–2458. https://doi.org/10.1021/jf011369q (2002).CAS  Article  PubMed  Google Scholar  Zielińska, E., BaRaniak, B. & Karaś, M. Antioxidant and anti-inflammatory activities of hydrolysates and peptide fractions obtained by enzymatic hydrolysis of selected heat-treated edible insects. Nutrients 9, 1–14 (2017). Google Scholar  Rahman, M. M., Byanju, B., Grewell, D. & Lamsal, B. P. High-power sonication of soy proteins: Hydroxyl radicals and their effects on protein structure. Ultrason. Sonochem. 64, 105019 (2020).CAS  PubMed  Google Scholar  More

Wu, X. et al. Effects of bio-organic fertiliser fortified by Bacillus cereus QJ-1 on tobacco bacterial wilt control and soil quality improvement. Biocontrol Sci. Technol. 30, 351–369 (2020).
Google Scholar 
Hu, W. et al. Flue-cured tobacco (Nicotiana tabacum L.) leaf quality can be improved by grafting with potassium-efficient rootstock. Field Crop. Res. 274, 108305 (2021).
Google Scholar 
Wu, X. et al. Bioaugmentation of Bacillus amyloliquefaciens-Bacillus kochii co-cultivation to improve sensory quality of flue-cured tobacco. Arch. Microbiol. 203, 5723–5733 (2021).CAS 
PubMed 

Google Scholar 
Jiang, C. et al. Optimal lime application rates for ameliorating acidic soils and improving the yield and quality of tobacco leaves. Appl. Ecol. Environ. Res. 18, 5411–5423 (2020).
Google Scholar 
Yin, Q. et al. Investigation of associations between rhizosphere microorganisms and the chemical composition of flue-cured tobacco leaves using canonical correlation analysis. Commun. Soil Sci. Plant 44, 1524–1539 (2013).CAS 

Google Scholar 
Shen, H. et al. Promotion of lateral root growth and leaf quality of flue-cured tobacco by the combined application of humic acids and npk chemical fertilizers. Exp. Agric. 53, 59–70 (2017).CAS 

Google Scholar 
Hu, W. Q. et al. Grafting alleviates potassium stress and improves growth in tobacco. BMC Plant Biol. 19, 130 (2019).PubMed 
PubMed Central 

Google Scholar 
Zhong, J. Study of K+ uptake kinetics of flue-cured tobacco in K+-enriched and conventional tobacco genotypes. J. Plant Nutr. 42(7), 1–7 (2019).
Google Scholar 
Tang, Z. et al. Climatic factors determine the yield and quality of Honghe flue-cured tobacco. Sci. Rep. 10, 19868 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Yan, S. et al. Correlation between soil microbial communities and tobacco aroma in the presence of different fertilizers. Ind. Crop. Prod. 151, 112454 (2020).CAS 

Google Scholar 
Tabaxi, I. Effect of organic fertilization on quality and yield of oriental tobacco (Nicotiana tabacum L.) under Mediterranean conditions. Asian J. Agric. Biol. https://doi.org/10.35495/ajab.2020.05.274 (2021).Article 

Google Scholar 
Chen, Y. L. et al. Distinct microbial communities in the active and permafrost layers on the Tibetan Plateau. Mol. Ecol. 26, 6608–6620 (2017).CAS 
PubMed 

Google Scholar 
Wagg, C. et al. Soil biodiversity and soil community composition determine ecosystem multifunctionality. Proc. Natl. Acad. Sci. U.S.A. 111, 5266–5270 (2014).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
French, E. et al. Emerging strategies for precision microbiome management in diverse agroecosystems. Nat. Plants 7, 256–267 (2021).PubMed 

Google Scholar 
Cui, Y. et al. Diversity patterns of the rhizosphere and bulk soil microbial communities along an altitudinal gradient in an alpine ecosystem of the eastern Tibetan Plateau. Geoderma 338, 118–127 (2019).ADS 
CAS 

Google Scholar 
Zheng, J. et al. The effects of tetracycline residues on the microbial community structure of tobacco soil in pot experiment. Sci. Rep. 10, 8804 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Zhang, J. Effects of tobacco planting systems on rates of soil N transformation and soil microbial community. Int. J. Agric. Biol. 19, 992–998 (2017).CAS 

Google Scholar 
Yang, Y. et al. Metagenomic insights into effects of wheat straw compost fertiliser application on microbial community composition and function in tobacco rhizosphere soil. Sci. Rep. 9, 6168 (2019).ADS 
PubMed 
PubMed Central 

Google Scholar 
Wang, S. et al. Response of soil fungal communities to continuous cropping of flue-cured tobacco. Sci. Rep. 10, 19911 (2020).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Finlay, B. J. Global dispersal of free-living microbial eukaryote species. Science 296, 1061–1063 (2002).ADS 
CAS 
PubMed 

Google Scholar 
Ehrmann, J. & Ritz, K. Plant: soil interactions in temperate multi-cropping production systems. Plant Soil 376, 1–29 (2013).
Google Scholar 
Liu, H. et al. Response of soil fungal community structure to long-term continuous soybean cropping. Front. Microbiol. 9, 3316 (2018).PubMed 

Google Scholar 
Gao, Z. et al. Effects of continuous cropping of sweet potato on the fungal community structure in rhizospheric soil. Front. Microbiol. 10, 2269 (2019).PubMed 
PubMed Central 

Google Scholar 
Li, J. Analysis on the method selection of tobacco disease control. South China Agric. 15, 49–50 (2021).
Google Scholar 
Dai, C. et al. Comprehensive evaluation of soil fertility status of tobacco-planting district in Bijie area. Acta Agric. Jiangxi 23, 9–11 (2011).
Google Scholar 
Wang, Z. et al. Time-course relationship between environmental factors and microbial diversity in tobacco soil. Sci. Rep. 9, 19969 (2019).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Wang, X. et al. Study on the primary chemical components, sensory quality of flue-cured tobacco and their correlativity in Bijie. J. Henan Agric. Sci. 41, 58–61 (2012).
Google Scholar 
Zhang, S. et al. Analysis of variation characteristics and coordination of conventional chemical components of flue-cured tobacco in Youyang County, Chongqing City. Acta Agric. Jiangxi 32, 75–86 (2020).
Google Scholar 
Lakatos, L. et al. The influence of meteorological variables on sour cherry quality parameters. In Vi International Cherry Symposium (Int Soc Horticultural Science, Leuven 1), Vol. 1020, 287–292 (2014).Zhao, Z. et al. Why does potassium concentration in flue-cured tobacco leaves decrease after apex excision? Field Crop. Res. 116, 86–91 (2010).
Google Scholar 
Travlos, I. S. et al. Green manure and pendimethalin impact on oriental sun-cured tobacco. Agron. J. 106, 1225–1230 (2014).
Google Scholar 
Bilalis, D. et al. Effect of organic fertilization on soil characteristics, yield and quality of Virginia Tobacco in Mediterranean area. Emir. J. Food Agric. https://doi.org/10.9755/ejfa.2020.v32.i8.2138 (2020).Article 

Google Scholar 
Henry, J. B., Vann, M. C. & Lewis, R. S. Agronomic practices affecting nicotine concentration in flue-cured tobacco: A review. Agron. J. 111, 3067–3075 (2019).CAS 

Google Scholar 
Lamarre, M. & Payette, S. Influence of nitrogen-fertilization on Quebec flue-cured tobacco production. Can. J. Plant Sci. 72, 411–419 (1992).CAS 

Google Scholar 
Zhang, L. et al. Dynamic changes of nutrients in tobacco-planting soils in Bijie City during 2014–2016. Guizhou Agric. Sci. 45, 51–55 (2017).CAS 

Google Scholar 
Lisuma, J. B., Mbega, E. R. & Ndakidemi, P. A. Dynamics of nicotine across the soil–tobacco plant interface is dependent on agro-ecology, nitrogen source, and rooting depth. Rhizosphere 12, 100175 (2019).
Google Scholar 
Cosme, M. & Wurst, S. Interactions between arbuscular mycorrhizal fungi, rhizobacteria, soil phosphorus and plant cytokinin deficiency change the root morphology, yield and quality of tobacco. Soil Biol. Biochem. 57, 436–443 (2013).CAS 

Google Scholar 
Chandanie, W. A., Kubota, M. & Hyakumachi, M. Interactions between the arbuscular mycorrhizal fungus Glomus mosseae and plant growth-promoting fungi and their significance for enhancing plant growth and suppressing damping-off of cucumber (Cucumis sativus L.). Appl. Soil Ecol. 41, 336–341 (2009).
Google Scholar 
Liu, D. et al. Geographic distance and soil microbial biomass carbon drive biogeographical distribution of fungal communities in Chinese Loess Plateau soils. Sci. Total Environ. 660, 1058–1069 (2019).ADS 
CAS 
PubMed 

Google Scholar 
Li, S. & Wu, F. Diversity and co-occurrence patterns of soil bacterial and fungal communities in seven intercropping systems. Front. Microbiol. https://doi.org/10.3389/fmicb.2018.01521 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Li, H. et al. Chaetosemins A-E, new chromones isolated from an Ascomycete Chaetomium seminudum and their biological activities. RSC Adv. 5, 29185–29192 (2015).ADS 
CAS 

Google Scholar 
Gorte, O., Kugel, M. & Ochsenreither, K. Optimization of carbon source efficiency for lipid production with the oleaginous yeast Saitozyma podzolica DSM 27192 applying automated continuous feeding. Biotechnol. Biofuels 13, 181 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Zhang, J. et al. Biochar applied to consolidated land increased the quality of an acid surface soil and tobacco crop in Southern China. J. Soil. Sediment. 20, 3091–3102 (2019).
Google Scholar 
Zong, J. et al. Effect of two drying methods on chemical transformations in flue-cured tobacco. Dry Technol. https://doi.org/10.1080/07373937.2020.1779287 (2020).Article 

Google Scholar 
Ismail, E. et al. Evaluation of in vitro antifungal activity of potassium bicarbonate on Rhizoctonia solani AG 4 HG-I, Sclerotinia sclerotiorum and Trichoderma sp.. Afr. J. Biotechnol. 10, 8605–8612 (2011).
Google Scholar 
d’Aquino, L. et al. Effect of some rare earth elements on the growth and lanthanide accumulation in different Trichoderma strains. Soil Biol. Biochem. 41, 2406–2413 (2009).CAS 

Google Scholar 
Bijie Municipal Government. Statistical Yearbooks of Bijie City (2016)He, R., Liu, S. & Liu, Y. Application of SD model in analyzing the cultivated land carrying capacity: A case study in Bijie Prefecture, Guizhou Province, China. Procedia Environ. Sci. 10, 1985–1991 (2011).
Google Scholar 
Wang, M. et al. Spatial variation and fractionation of fluoride in tobacco-planted soils and leaf fluoride concentration in tobacco in Bijie City, Southwest China. Environ. Sci. Pollut. Res. 28, 26112–26123 (2021).CAS 

Google Scholar 
Wang, J. T. et al. Altitudinal distribution patterns of soil bacterial and archaeal communities along Mt. Shegyla on the Tibetan Plateau. Microb. Ecol. 69, 135–145 (2015).PubMed 

Google Scholar 
Zhang, Q. et al. Soil available phosphorus content drives the spatial distribution of archaeal communities along elevation in acidic terrace paddy soils. Sci. Total Environ. 658, 723–731 (2019).ADS 
CAS 
PubMed 

Google Scholar 
Cui, Q. et al. Sulfur application improved leaf yield and quality of flue-cured tobacco by maintaining soil sulfur balance. Int. J. Agric. Biol. 23, 357–363 (2020).CAS 

Google Scholar 
Cao, X. et al. Distribution, availability and translocation of heavy metals in soil-oilseed rape (Brassica napus L.) system related to soil properties. Environ. Pollut. 252, 733–741 (2019).CAS 
PubMed 

Google Scholar 
Wang, C. et al. Prevalence of antibiotic resistance genes and bacterial pathogens along the soil-mangrove root continuum. J. Hazard. Mater. 408, 124985 (2021).CAS 
PubMed 

Google Scholar 
Magoc, T. & Salzberg, S. L. FLASH: Fast length adjustment of short reads to improve genome assemblies. Bioinformatics 27, 2957–2963 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: A flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014).CAS 
PubMed 
PubMed Central 

Google Scholar 
Edgar, R. C. et al. UCHIME improves sensitivity and speed of chimera detection. Bioinformatics 27, 2194–2200 (2011).CAS 
PubMed 
PubMed Central 

Google Scholar 
Edgar, R. C. UPARSE: Highly accurate OTU sequences from microbial amplicon reads. Nat. Methods 10, 996–998 (2013).CAS 
PubMed 

Google Scholar 
Bokulich, N. A. et al. Quality-filtering vastly improves diversity estimates from Illumina amplicon sequencing. Nat. Methods 10, 57–59 (2013).CAS 
PubMed 

Google Scholar 
Wang, Q. et al. Naive Bayesian classifier for rapid assignment of rRNA sequences into the new bacterial taxonomy. Appl. Environ. Microbiol. 73, 5261–5267 (2007).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Koljalg, U. et al. Towards a unified paradigm for sequence-based identification of fungi. Mol. Ecol. 22, 5271–5277 (2013).CAS 
PubMed 

Google Scholar 
Bolyen, E. et al. Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2. Nat. Biotechnol. 37, 852–857 (2019).CAS 
PubMed 
PubMed Central 

Google Scholar 
Lindstrom, E. S. et al. Distribution of typical freshwater bacterial groups is associated with pH, temperature, and lake water retention time. Appl. Environ. Microbiol. 71, 8201–8206 (2005).ADS 
PubMed 
PubMed Central 

Google Scholar  More

Phylogenetic analysesA total of 21 sequences were newly generated and deposited in GenBank (Supplementary Table 1). The concatenated sequence alignment of the three loci comprised 100 sequences (38 for ITS, 30 for rpb2 and 32 for mtSSU) from 43 collections (Supplementary Table 1). The alignment was 2,004 characters long, including gaps. Multi-locus trees generated from ML and BI analyses showed similar topologies without any supported topological conflict. The multi-locus phylogeny (Fig. 1) confirmed placement of all Thai collections within the well-supported R. subsect. Amoeninae (ML = 99, BI = 1.0). Five collections from northeastern Thailand and two collections from northern Thailand form two strongly supported clades and are described below as the new species R. bellissima sp. nov. and R. luteonana sp. nov. The new species are not resolved as sister. The first species, R. bellissima, is strongly supported as sister to a clade of Australian sequestrate species that includes R. variispora T. Lebel and an undescribed Russula sp. labeled as Macowanites sp. The Indian species R. intervenosa S. Paloi, A.K. Dutta & K. Acharya is placed as sister to them with bootstrap support of 77. The second species, R. luteonana, is placed with moderate support as sister to the sequestrate European species R. andaluciana T.F. Elliott & Trappe.Figure 1ML phylogenetic tree inferred from the three-gene dataset (ITS, rpb2, mtSSU) of Russula subsection Amoeninae species, using ML and BI analyses. Three members of R. subg. Heterophyllidiae are used as outgroup. Species in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageThe ITS tree (Fig. 2) shows a similar topology and relationships for the studied specimens. In addition, R. intervenosa received good support (ML = 84, BI = 0.99) as sister to the clade of R. bellissima and R. variispora. Five additional ITS sequences that are grouped with strong support within R. bellissima species clade were recovered, three from Thailand, one from Laos, and one from Singapore. We did not recover any other Amoeninae ITS sequences from Thailand.Figure 2ML phylogenetic tree inferred from the ITS region of Russula subsection Amoeninae species and allied groups, using ML and BI methods. Samples in boldface are new species in this study. Bootstrap support values (BS ≥ 50%) and posterior probabilities (PP ≥ 0.90) are shown at the supported branches.Full size imageTaxonomy
Russula bellissima Manz & F. Hampe sp. nov.
Mycobank: MB 840549Holotype THAILAND, Theong district, Chiang Rai, 19°36′45”N 100°4′00”E, alt. 500 m, dry dipterocarpus forest in small groups on loamy soil, 12 July 2012, F. Hampe (Holotype: GENT FH 12-127; Isotype: MFLU12-0619).Etymology ’bellus’ = latin for beautiful, pretty, lovely; ’bellissima’ = the most beautiful. Resembling the species Russula bella which is also belonging to Russula subsection Amoeninae.Diagnosis Pileus small to medium-sized; cuticle dry, smooth, matt and pruinose, red; stipe white or with a red flush; spore ornamentation of moderately distant to dense amyloid spines or warts, frequently fused into short crests or even long wings; suprahilar spot inamyloid; hymenial cystidia and pileocystidia absent.Pileus (Fig. 3) small to medium sized, 10–50 mm diam., young hemispherical or convex, becoming plane and depressed at the centre; margin first even, when old distinctly tuberculate-striate up to 10 mm from the margin, often radially cracking; cuticle hardly peeling, radially disrupted into small patches, pruinose when young, later dry, smooth, matt and pruinose in the centre, colour near the margin when young varnish red (9C8), later red to coral red (9B6-7); near the centre deep red, blood red, dark red (10C7-8), raspberry red (10D7), strawberry red (10D8) or purple brown (10E-F8). Lamellae: 3–5 mm deep, thin, moderately dense, 6–8 at 1 cm near the pileus margin, adnexed, white, slightly anastomosing at the base; lamellulae absent, occasionally forked near the stipe; edges concolorous, entire but pruinose under lens. Stipe: 10–30 × 3–7 mm, usually narrowed towards the base, sometimes cylindrical, surface smooth, white and mainly with a distinct pastel red to red flush, occasionally completely white or sometimes also almost completely red, interior stuffed. Context: white, fragile, unchanging when damaged, reaction with guaiac after 5 s negative on both stipe and lamellae surfaces, reaction to FeSO4 and sulfovanillin negative; taste mild; odour inconspicuous. Spore print: not observed.Figure 3Basidiomata of Russula bellissima. (A) FH12-127 (Holotype). (B) FH12-158. Scale bar = 1 cm. Photos by Felix Hampe.Full size imageSpores (Figs. 4, 5) (6.9–)7.3–7.8–8.3(–8.9) × (6.1–)6.8–7.2–7.6(–8.4) µm, subglobose to broadly ellipsoid, Q = 1.01–1.1–1.2(–1.29); ornamentation of moderately distant [(4–)5–6(–7) in a 3 µm diam. circle] amyloid spines or warts, (1.1–)1.2–1.4–1.6(–1.7) µm high, fused or connected by fine line connections into often long crests or wings, [(0–)1–3(–4) fusions and the same number of line connections in a 3 µm diam. circle], crests and wings frequently branched and occasionally form closed loops, isolated elements dispersed, edge of crests and wings irregularly wavy; suprahilar spot moderately large, inamyloid. Basidia: (30.5–)34.5–44.1–53.5(–65.0) × (10.5–)11.5–12.6–14.0(–16.0) µm, broadly clavate or obpyriform, 4-spored; basidiola cylindrical, ellipsoid or broadly clavate, ca. 5–10 µm wide. Hymenial cystidia on lamellae sides: absent. Lamellae edges: covered by densely arranged or fasciculate marginal cells. Marginal cells: (27.0–)38.5–46.4–54.5(–61.0) × (5.0–)5.5–6.7–7.5(–9.0) µm; subulate or narrowly lageniform, apically attenuated and constricted to ca. 1–2 µm, sometimes slightly moniliform or flexuous. Pileipellis: (Fig. 6) orthochromatic in Cresyl Blue, gradually passing to the underlying context, 200–300 µm deep; suprapellis 60–130 µm deep, composed of erect or ascending hyphal terminations forming a dry trichoderm, well delimited from 140 to 210 µm deep subpellis composed of horizontally oriented, strongly gelatinized narrow hyphae. Subpellis not well delimited from the underlying context, elongate hyphae gradually changing to sphaerocytes. Acid- resistant incrustations: absent. Hyphal terminations near the pileus margin: composed of long apically attenuated terminal cell and a chain of 1–4 ovoid to barrel shaped, short unbranched cells with one distinctly longer apical cell; constricted on septa, usually not flexuous, oriented towards the pileus surface, usually thin-walled, sometimes slightly thick-walled (up to 1 µm thick); terminal cells mainly subulate or lageniform, apically attenuated and acute, measuring (19–)27.5–38.3–49.0(–66.5) × (3.3–)4.5–5.8–7.0(–9.0) µm, rarely with a forked apex, mixed with dispersed, cylindrical or ellipsoid, distinctly shorter, obtuse terminal cells measuring (7.5–)11.5–17.8–29.5(–42.5) × (3.0–)4.0–4.5–5.0 µm; subterminal cells measuring (4.5–)5.5–8.3–11.5(–16.0) × 4.5–5.3–6.0(–7.0) µm. Hyphal terminations near the pileus centre: similar in shape and also with a mixture of long acute and short obtuse terminal cells, acute ones measuring (12.0–)22.0–35.2–48.5(–79.0) × (2.5–)3.5–4.9–6.5(–8.0) µm, obtuse ones more frequent, measuring (6.5–)8.5–12.0–15.5(–22.0) × (3.5–)4.0–4.9–6.0(–7.5) µm. Primordial hyphae or pileocystidia: absent. Cystidioid hyphae and oleipherous hyphae not observed.Figure 4Hymenial elements of Russula bellissima (holotype, FH 12-127). (A) Basidia and basidiolae. (B) Marginal cells. (C) Spores as seen in Melzer’s reagent. Scale bar = 10 µm, but only 5 µm for spores.Full size imageFigure 5Scanning electron microscope photo of spore ornamentation. Russula bellissima (holotype, FH 12-127). Scale bar = 2 μm.Full size imageFigure 6Elements of the pileipellis of Russula bellissima (holotype, FH 12-127). (A) Hyphal terminations near the pileus margin. (B) Hyphal terminations near the pileus centre. Scale bar = 10 μm.Full size imageAdditional material studied THAILAND, Chiang Mai Province, Mae On District, about 3 km from Tharnthong lodges, 18° 51′ 55″ N 99° 17′ 23″ E, alt. 725 m, Dipterocarpaceae dominated forest with the presence of some Castanopsis trees, in small groups on loamy soil, 17 July 2012, F. Hampe (GENT FH 12-158, duplicate: MFLU12-0648).Note Russula bellissima is a small species with a bright red pileus and pink colour on the stipe. This colour is distinctive and resembles North American R. mariae, Indian R. intervenosa and Asian R. bella. It is very unlikely that the distribution of any European or North American species is overlapping with the Thai species. However, little is known about the distributional ranges and the ecological niches of other Asian Russula species. Therefore discussing the morphological distinguishing characters between Asian species and R. bellissima is more relevant. Russula bellissima is not closely related to R. bella and it differs from this species by larger spores with a more prominent spore ornamentation, absence of hymenial cystidia on lamellae sides, and subterminally short, ellipsoid cells in the suprapellis arranged in unbranched chains of up to four7. The Thai species resembles and is closely related to the Indian R. intervenosa, but it has a more prominent spore ornamentation, hymenial cystidia (on lamellae sides) are absent, and hyphal terminations in the pileipellis are wider22.
Russula luteonana M. Pobkwamsuk & K. Wisitrassameewong sp. nov.
Mycobank: MB 840550Holotype: THAILAND, Amnat Charoen province, Hua Taphan district, Junction near Watbochaneng , dry dipterocarp forest, alt. 145 m, 15° 41′ 28″ N 104° 31′ 41″ E, 13 July 2016, Thitiya Boonpratuang, Rattaket Choeyklin, Prapapan Sawhasan, Maneerat Pobkwamsuk, Pattrachai Juthamas, Nattawut Wiriyathanawudhiwong, Patcharee Patangwesa (BBH41120).Etymology ‘Luteolus’ = yellow colour, ‘Nanus’ = small. Refer to pileus color and size of the species.Diagnosis Pileus medium-sized, dry, usually yellow, spores with subreticulate amyloid ornamentation and inamyloid suprahilar spot, hymenial cystidia on lamellae sides large, lamellae edges with combination of subulate, clavate and pyriform marginal cells.Pileus (Fig. 7) medium-sized, 28‒53 mm diam., plano-convex with depressed centre, infundibuliform when mature; margin striated and radially cracking in dry condition; cuticle dry, peeling to almost ½ of radius, smooth to minutely wrinkled, dull in dry condition, color very variable, some collections pale cream and with darker pale brownish-yellow centre, other yellow brownish and with darker orange-brown centre, sometimes also bright red-brown and with discolored centre, always with rusty-brown spots especially when near the centre. Lamellae: 3‒5 mm deep, moderately distant, intervenose, forking near the stipe, white to cream, edges even, concolorous. Stipe: 26‒40 × 6‒9 mm, cylindrical or narrowed at the base, surface dry, longitudinally wrinkled, white, turning brown when bruised. Context: 2‒4 mm in at the half pileus radius, soft, solid, becoming partially hollow when mature, white, unchanging when cut. Taste mild; odour rather strong, fishy. Spore print: not observed.Figure 7Basidiomata of Russula luteonana. (A) BBH41120 (Holotype). (B) BBH41121. (C) BBH41122. (D) BBH42510. Scale bar = 1 cm. Photos by Thitiya Boonpratuang.Full size imageSpores (Figs. 8, 9) (7.4‒)8.1‒8.6‒9(‒10.1) × (6.1‒)7.4‒7.5‒7.9(‒9.1) μm, subglobose to broadly ellipsoid, Q = (1.03‒)1.09‒1.15‒1.20(‒1.30), ornamentation of moderately distant, obtuse, (0.7‒)1.1‒1.3‒1.5(‒1.9) μm high spines, connected by abundant line connections [(0‒)3‒6(‒8) in in a 3 µm diam. circle], branched, forming an incomplete reticulum, crest irregularly wavy and occasionally fused [(0‒)1‒2(‒5) fusions in the circle], isolated elements rare; suprahilar spot inamyloid. Basidia: (29‒)34.5‒39.1‒44(‒51.5) × (10‒)12‒13.2‒14.5(‒16.5) μm, clavate, 4-spored, rarely 2-spored, basidiola subcylindrical to subclavate, (25.5‒)30‒35.4‒41(‒47) × (9‒)11‒12.2‒14 (‒16) μm. Hymenial cystidia on lamellae sides: usually protruding over other elements of hymenium, widely dispersed ( More

Lincoln, G. A. Teeth, horns and antlers: the weapons of sex. In The Differences between the Sexes (eds R. V. Short & E. Balaban) 131–158 (Cambridge Univ. Press, 1994).Lundrigan, B. Morphology of horns and fighting behavior in the family bovidae. J. Mammal. 77, 462–475 (1996).Article 

Google Scholar 
Bro-Jorgensen, J. The intensity of sexual selection predicts weapon size in male bovids. Evolution 61, 1316–1326 (2007).Article 

Google Scholar 
Plard, F., Bonenfant, C. & Gaillard, J. M. Revisiting the allometry of antlers among deer species: male-male sexual competition as a driver. Oikos 120, 601–606 (2011).Article 

Google Scholar 
Okada, K. & Miyatake, T. Sexual dimorphism in mandibles and male aggressive behavior in the presence and absence of females in the beetle Librodor japonicus (Coleoptera: Nitidulidae). Ann. Entomol. Soc. Am. 97, 1342–1346 (2004).Article 

Google Scholar 
Emlen, D. J., Marangelo, J., Ball, B. & Cunningham, C. W. Diversity in the weapons of sexual selection: Horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution 59, 1060–1084 (2005).CAS 
Article 

Google Scholar 
Pomfret, J. C. & Knell, R. J. Sexual selection and horn allometry in the dung beetle Euoniticellus intermedius. Anim. Behav. 71, 567–576 (2006).Article 

Google Scholar 
McCullough, E. L., Weingarden, P. R. & Emlen, D. J. Costs of elaborate weapons in a rhinoceros beetle: how difficult is it to fly with a big horn?. Behav. Ecol. 23, 1042–1048 (2012).Article 

Google Scholar 
David, P., Bjorksten, T., Fowler, K. & Pomiankowski, A. Condition-dependent signalling of genetic variation in stalk-eyes flies. Nature 406, 186–188 (2000).ADS 
CAS 
Article 

Google Scholar 
Baker, R. H. & Wilkinson, G. S. Phylogenetic analysis of sexual dimorphism and eye-span allometry in stalk-eyed flies (Diopsidae). Evolution 55, 1373–1385 (2001).CAS 
Article 

Google Scholar 
Stankowich, T. Armed and dangerous: predicting the presence and function of defensive weaponry in mammals. Adapt. Behav. 20, 32–43 (2012).Article 

Google Scholar 
Hashimoto, K. & Hayashi, F. Structure and function of the large pronotal horn of the sand-living anthicid beetle Mecynotarsus tenuipes. Entomol. Sci. 15, 274–279 (2012).Article 

Google Scholar 
Hayashi, M. & Ohba, S. Y. Mouth morphology of the diving beetle Hyphydrus japonicus (Dytiscidae: Hydroporinae) is specialized for predation on seed shrimps. Biol. J. Linn. Soc. 125, 315–320 (2018).Article 

Google Scholar 
Stocker, R. F. The organization of the chemosensory system in Drosophila melanogaster: a review. Cell Tissue Res. 275, 3–26 (1994).CAS 
Article 

Google Scholar 
Dweck, H. K. M. Antennal sensory receptors of Pteromalus puparum female (Hymenoptera: Pteromalidae), a gregarious pupal endoparasitoid of Pieris rapae. Micron 40, 769–774 (2009).Article 

Google Scholar 
Crespo, J. G. A review of chemosensation and related behavior in aquatic insects. J. Insect Sci. 11, 1–39 (2011).Article 

Google Scholar 
Stoffolano, J. G. Jr., Rice, M. & Murphy, W. L. The importance of antennal mechanosensilla of Sepedon fuscipennis (Diptera: Sciomyzidae). Can. Entomol. 145, 265–272 (2013).Article 

Google Scholar 
Gabel, B. et al. Floral volatiles of Tanacetum vulgare L. attractive to Lobesia botrana Den. et Schiff. females. J. Chem. Ecol. 18, 693–701 (1992).CAS 
Article 

Google Scholar 
Fox, H. Barbels and barbel-like tentacular structures in sub-mammalian vertebrates: a review. Hydrobiologia 403, 153–193 (1999).Article 

Google Scholar 
Plepys, D., Ibarra, F., Francke, W. & Lofstedt, C. Odour-mediated nectar foraging in the silver Y moth, Autographa gamma (Lepidoptera: Noctuidae): behavioural and electrophysiological responses to floral volatiles. Oikos 99, 75–82 (2002).CAS 
Article 

Google Scholar 
Stankowich, T. & Caro, T. Evolution of weaponry in female bovids. Proc. R. Soc. Lond. Ser. B Biol. Sci. 276, 4329–4334 (2009).Bergmann, P. J. & Berk, C. P. The Evolution of Positive Allometry of Weaponry in Horned Lizards (Phrynosoma). Evol. Biol. 39, 311–323 (2012).Article 

Google Scholar 
Damman, H. The osmaterial glands of the swallowtail butterfly Eurytide marcellus as a defense against natural enemies. Ecol. Entomol. 11, 261–265 (1986).Article 

Google Scholar 
Berenbaum, M. R., Moreno, B. & Green, E. Soldier bug predation on swallowtail caterpillars (Lepidoptera, Papilionidae): circumvention of defensive chemistry. J. Insect Behav. 5, 547–553 (1992).Article 

Google Scholar 
Juma, G. et al. Distribution of chemo- and mechanoreceptors on the antennae and maxillae of Busseola fusca larvae. Entomol. Exp. Appl. 128, 93–98 (2008).Article 

Google Scholar 
Liu, Z., Hua, B.-Z. & Liu, L. Ultrastructure of the sensilla on larval antennae and mouthparts in the peach fruit moth, Carposina sasakii Matsumura (Lepidoptera: Carposinidae). Micron 42, 478–483 (2011).Article 

Google Scholar 
Kandori, I., Tsuchihara, K., Suzuki, T. A., Yokoi, T. & Papaj, D. R. Long frontal projections help Battus philenor (Lepidoptera: Papilionidae) larvae find host plants. PLoS ONE 10, e0131596 (2015).Article 

Google Scholar 
Greeney, H. F., Dyer, L. A. & Smilanich, A. M. Feeding by lepidopteran larvae is dangerous: A review of caterpillars’ chemical, physiological, morphological, and behavioral defenses against natural enemies. ISJ Invert. Surviv. J. 9, 7–34 (2012).
Google Scholar 
Sugiura, S. Predators as drivers of insect defenses. Entomol. Sci. 23, 316–337 (2020).Article 

Google Scholar 
Martin, W. R. & Nordlund, D. A. Ovipositional behavior of the parasitoid Palexorista laxa (Diptera, Tachinidae) on Heliothis zea (Lepidoptera, Noctuidae) larvae. J. Entomol. Sci. 24, 460–464 (1989).Article 

Google Scholar 
Constantino, L. M. Notes on Haetera from Colombia, with description of the immature stages of Haetera piera (Lepidoptera:Nymphalidae: Satyrinae). Trop. Lepid. 4(1), 13–15 (1993).
Google Scholar 
Devries, P. J., Kitching, I. J. & Vanewright, R. I. The systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera). Syst. Entomol. 10, 11–32. https://doi.org/10.1111/j.1365-3113.1985.tb00561.x (1985).Article 

Google Scholar 
Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Biology and external morphology of immature stages of Memphis appias (Hubner) (Lepidoptera: Nymphalidae: Charaxinae). Zootaxa, 21–32 (2010).Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Biology and external morphology of the immature stages of the butterfly Callicore pygas eucale, with comments on the taxonomy of the genus Callicore (Nymphalidae: Biblidinae). J. Insect Sci. 14, doi:https://doi.org/10.1093/jis/14.1.91 (2014).Dias, F. M. S., Casagrande, M. M. & Mielke, O. H. H. Immature stages of the turquoise-banded shoemaker Archaeoprepona amphimachus pseudomeander (Fruhstorfer, 1906) and a comparative review of the Preponini (Lepidoptera: Nymphalidae). Aust. Entomol. 58, 451–462. https://doi.org/10.1111/aen.12339 (2019).Article 

Google Scholar 
Dias, F. M. S., de Oliveira-Neto, J. F., Casagrande, M. M. & Mielke, O. H. H. External morphology of immature stages of Zaretis strigosus (Gmelin) and Siderone galanthis catarina Dottax and Pierre comb. nov., with taxonomic notes on Siderone (Lepidoptera: Nymphalidae: Charaxinae). Rev. Bras. Entomol. 59, 307–319, doi:https://doi.org/10.1016/j.rbe.2015.07.007 (2015).Dias, F. M. S. et al. An integrative approach elucidates the systematics of Sea Hayward and Cybdelis Boisduval (Lepidoptera: Nymphalidae: Biblidinae). Syst. Entomol. 44, 226–250. https://doi.org/10.1111/syen.12327 (2019).Article 

Google Scholar 
Freitas, A. V. L., Barbosa, E. P. & Marin, M. A. Immature Stages and Natural History of the Neotropical Satyrine Pareuptychia ocirrhoe Interjecta (Nymphalidae: Euptychiina). J. Lepid. Soc. 70, 271–276. https://doi.org/10.18473/lepi.70i4.a4 (2016).Article 

Google Scholar 
Freitas, A. V. L., Kaminski, L. A., Mielke, O. H. H., Barbosa, E. P. & Silva-Brandao, K. L. A new species of Yphthimoides (Lepidoptera: Nymphalidae: Satyrinae) from the southern Atlantic forest region. Zootaxa, 31–44 (2012).Furtado, E. & Campos-Neto, F. C. Caligopsis seleucida (Hewitson) and its immature stages (Lepidoptera, Nymphalidae, Brassolinae). Rev. Bras. Zool. 21(3), 593–597 (2004).Article 

Google Scholar 
Greeney, H. F. et al. The early stages and natural history of Antirrhea adoptiva porphyrosticta (Watkins, 1928) in eastern Ecuador (Lepidoptera: Nymphalidae: Morphinae). J. Insect Sci. 9 (2009).Greeney, H. F. et al. Early stages and natural history of Perisama oppelii (Nymphalidae, Lepidoptera) in eastern Ecuador. Kempffiana 6(1), 16–30 (2010).
Google Scholar 
Greeney, H. F., Dyer, L. A. & Pyrcz, T. W. First description of the early stage biology of the genus Mygona: The natural history of the satyrine butterfly, Mygona irmina in eastern Ecuador. J. Insect Sci. 11, doi:https://doi.org/10.1673/031.011.0105 (2011).Greeney, H. F., Pyrcz, T. W., DeVries, P. J. & Dyer, L. A. The early stages of Pedaliodes poesia (Hewitson, 1862) in eastern Ecuador (Lepidoptera: Satyrinae: Pronophilina). J. Insect Sci. 9 (2009).Greeney, H. F., Whitfield, J. B., Stireman, J. O., Penz, C. M. & Dyer, L. A. Natural history of Eryphanis greeneyi (Lepidoptera: Nymphalidae) and its enemies, with a description of a new species of Braconid parasitoid and notes on its Tachinid parasitoid. Ann. Entomol. Soc. Am. 104, 1078–1090. https://doi.org/10.1603/an10064 (2011).Article 

Google Scholar 
Kaminski, L. A. & Freitas, A. V. L. Immature stages of the butterfly Magneuptychia libye (L.) (Lepidoptera : Nymphalidae, Satyrinae). Neotrop. Entomol. 37, 169–172, doi:https://doi.org/10.1590/s1519-566×2008000200010 (2008).Lambkin, T. & Kendall, R. The status of Yoma algina (boisduval, 1832) & Y. sabina (cramer, 1780) (Lepidoptera: Nymphalidae: Nymphalinae) in Australia. Aust. Entomol. 43 (4), 211–234 (2016).Leite, L. A. R., Casagrande, M. M., Mielke, O. H. H. & Freitas, A. V. L. Immature stages of the Neotropical butterfly, Dynamine agacles agacles. J. Insect Sci. 12 (2012).Leite, L. A. R., Dias, F. M. S., Carneiro, E., Casagrande, M. M. & Mielke, O. H. H. Immature stages of the Neotropical cracker butterfly, Hamadryas epinome. J. Insect Sci. 12 (2012).Murillo, L. R. & Nishida, K. Life history of Manataria maculata (Lepidoptera : Satyrinae) from Costa Rica. Rev. Biol. Trop. 51, 463–469 (2003).PubMed 

Google Scholar 
Nakahara, S., Janzen, D. H., Hallwachs, W. & Espeland, M. Description of a new genus for Euptychia hilara (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 4012, 525-541, doi:https://doi.org/10.11646/zootaxa.4012.3.7 (2015).Penz, C. M., Freitas, A. V. L., Kaminski, L. A., Casagrande, M. M. & Devries, P. J. Adult and early-stage characters of Brassolini contain conflicting phylogenetic signal (Lepidoptera, Nymphalidae). Syst. Entomol. 38, 316–333. https://doi.org/10.1111/syen.12000 (2013).Article 

Google Scholar 
Pyrcz, T. W. et al. Uncovered diversity of a predominantly Andean butterfly clade in the Brazilian Atlantic forest: a revision of the genus Praepedaliodes Forster (Lepidoptera: Nymphalidae, Satyrinae, Satyrini). Neotrop. Entomol. 47, 211–255. https://doi.org/10.1007/s13744-017-0543-x (2018).CAS 
Article 
PubMed 

Google Scholar 
Shirai, L. T. et al. Natural history of Selenophanes cassiope guarany (Lepidoptera: Nymphalidae: Brassolini): an integrative approach, from molecules to ecology. Ann. Entomol. Soc. Am. 110, 145–159. https://doi.org/10.1093/aesa/saw068 (2017).Article 

Google Scholar 
Silva, P. L. et al. Immature Stages of the Brazilian Crescent Butterfly Ortilia liriope (Cramer) (Lepidoptera: Nymphalidae). Neotrop. Entomol. 40, 322–327. https://doi.org/10.1590/s1519-566×2011000300006 (2011).CAS 
Article 
PubMed 

Google Scholar 
Song-yun, L. Immature stages of Faunis aerope (Leech, 1890) (Lepidoptera, Nymphalidae). Atalanta 42, 221–222 (2011).
Google Scholar 
Steiner, H. Life history of Melanocyma faunula in Malaysia (Lepidoptera: Nymphalidae: Morphinae). Trop. Lepid. Res. 16, 23–26 (2005).
Google Scholar 
Velez, P. D., Montoya, H. H. V. & Wolff, M. Immature stages and natural history of the Andean butterfly Altinote ozomene (Nymphalidae: Heliconiinae: Acraeini). Zoologia 28, 593–602. https://doi.org/10.1590/s1984-46702011000500007 (2011).Article 

Google Scholar 
Wahlberg, N. et al. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proc. R. Soc. Lond. Ser. B Biol. Sci. 276, 4295–4302, doi:https://doi.org/10.1098/rspb.2009.1303 (2009).Willmott, K. R., Elias, M. & Sourakov, A. Two possible caterpillar mimicry complexes in neotropical Danaine butterflies (Lepidoptera: Nymphalidae). Ann. Entomol. Soc. Am. 104, 1108–1118. https://doi.org/10.1603/an10086 (2011).Article 

Google Scholar 
Willmott, K. R. & Freitas, A. V. L. Higher-level phylogeny of the Ithomiinae (Lepidoptera : Nymphalidae): classification, patterns of larval hostplant colonization and diversification. Cladistics 22, 297–368. https://doi.org/10.1111/j.1096-0031.2006.00108.x (2006).Article 
PubMed 

Google Scholar 
Zacca, T. et al. Revision of Godartiana Forster (Lepidoptera: Nymphalidae), with the description of a new species from northeastern Brazil. Aust. Entomol. 56, 169–190. https://doi.org/10.1111/aen.12223 (2017).Article 

Google Scholar 
Bossart, J.L., Fetzner Jr., J.F. & Rawlins, J.E. Ghana Butterfly Biodiversity Project website. https://www.invertebratezoology.org/GhanaBfly/default.asp (2007).Butterflies and Moths of North America project. Butterflies and Moths of North America website. https://www.butterfliesandmoths.org/ (2021).Dauphin, D. & Dauphin, J. The Rio Grande Valley’s Nature Site website. http://www.thedauphins.net (2021).Eeles, P. UK Butterflies website. https://www.ukbutterflies.co.uk/index.php. (2021).Florida Museum of Natural History. Florida Museum website. https://www.floridamuseum.ufl.edu/ (2021).Khew, S. K. et al. Butterflies of Singapore website. https://butterflycircle.blogspot.com/ (2021).Kunte, K., Sondhi, S. & Roy, P. Butterflies of India, v. 3.24. Indian Foundation for Butterflies website. https://www.ifoundbutterflies.org (2021).Miller, S. & Morrison, C. Parasitoid-Caterpillar-Plant Interactions in the Americas website. https://caterpillars.myspecies.info/ (2021).National Biodiversity Network Trust. iNaturalistUK website. https://uk.inaturalist.org/ (2021).Nature Picture Library Limited. Nature Picture Library website. https://www.naturepl.com/blog/ (2021).Project Noah Team. Project Noah website. https://www.projectnoah.org/ (2021).Shiraiwa, K. Pteron World. The encyclopedia website of the butterflies. https://www.pteron-world.com/index.html (2021).Wagner, W. Lepidoptera and Their Ecology website. http://www.pyrgus.de/ (2021).Wahlberg, N. & Peña, C. Nymphalidae.net. website. http://www.nymphalidae.net/ (2021).Wikimedia Foundation, Inc. Wikimedia Commons website. https://commons.wikimedia.org/ (2021).Matsuura, M. Social Wasps of Japan in Color. (in Japanese) (Hokkaido university press 2015).IBM SPSS. SPSS Base 25.0 User’s Guide. (SPSS Inc., 2017). More

Phyla Bdellovibrionota, Fusobacteriota, and Myxococcota were present in the green microbial mat but in negligible quantities in the brown mat. The unique phyla detected in the brown mat, but not in the green microbial mat, included Caldatribacteriota, Thermodesulfobacteriota, Dictyoglomota, Elusimicrobiota, Thermotogota, Candidatus Calescamantes, Fervidibacteria, Hydrothermae, GAL15 and TA06. The Candidatus Caldatribacterium (phyla Caldatribacteriota), earlier named OP9 was also detected in this work. Using single-cell and metagenome sequencing, data elucidated that Ca. Caldatribacterium conducts anaerobic sugar fermentation and exhibited diverse glycosyl hydrolases, including endoglucanase37.Cyanobacteria and Chloroflexota were the main identified phyla in the green microbial mat. Because the hot spring is almost stagnant, undisturbed, and the water surface temperature ( More

Aggregate size distributionAs hypothesized, the improved soil aggregation was observed under ICL, which is attributed to the presence of animals resulting in higher organic matter contents of total C and N fractions that can significantly enhance soil health over time32. Moreover, well-aggregated soils as observed under ICL ( > 4 mm) at site 1 and NE (2–4 mm) at site 2 have a greater potential of retaining their structure and may have higher macropores, which facilitate sustained root growth than soils with low aggregation such as under CNT (corn–soybean without grazing or CC) in this study. It also explains the significance of ICL systems with no-tillage and undisturbed grassland, where the formation of stable macroaggregates may occur as a result of incorporation of plant residues, stimulation of root exudates and increased biological activity. Furthermore, it was noticed that ICL system not only enhanced the macroaggregates but accentuated the presence of microaggregates due to persistent binding agents, which are critical in SOC protection against microbial decomposition. When integrating grazing livestock into crop rotation, soil aggregation is typically improved under moderate and controlled grazing than the high intensity grazing systems33. Compared to the long-term sites ( > 30 years), short-term site 4 did not result in discernible effects of grazing or CC on soil aggregation. However, within this short-term study, grazed pasture mix was able to enhance aggregation of 1–2 and 2–4 mm sized aggregates compared with oats, oats with CC, oats with CC and grazing. To observe the influence of CC and grazing on  > 4 mm or  4 mm) under ICL at site 1 resulted in 1.3–1.5 times significantly higher SOC concentration than NE and CNT. The greater concentration of SOC and N in ICL and NE is attributed to the lack of soil disturbance, crop residue retention, and rhizodeposition, which reduces macroaggregate turnover rate14. At site 3, NE enhanced aggregate-associated C and N concentrations and performed significantly better than both ICL and CNT treatments. The higher C and N accrual in the NE than ICL and CNT, especially at site 3, can be due to massive root systems, long-term establishment and absence of cultivation, which contributes to enhanced soil quality, while reducing nutrient vulnerability to loss by oxidation18,36. For short-term study at site 4, insignificant differences in aggregate-associated SOC suggested that longer study period of at least  > 5 years is required for SOC to respond to grazing and cover crop management. The higher total N under ICL and NE can also be due to the presence of legumes, and brassicas in CC, which are effective at recycling N and may have helped in scavenging N.An overall increase in C and N cycling under ICL and NE systems has been attributed to ingested pasture being converted into urine and manure. Under these systems, livestock catalyze nutrient cycling by breakdown of complex plant molecules, greater soil incorporation and decomposition of plant residues and soil organic matter, which can maintain or even improve soil fertility by production of organic acids such as fulvic and humic acids6,8,19. Moreover, grazing stimulates the carbohydrate exudation from grass roots, which is mostly composed of polysaccharides, a C-O alkyl source37. The enhanced C concentration under ICL and NE can also be associated with higher MWD. Integrated system cool-season pasture and winter CC had significantly higher total C and N than the non-integrated continuous corn in previous study6. The results from another integrated system study7 showed that soybean and oat-Italian ryegrass CC increased total C (1.16 Mg ha−1 yr−1) and N stocks (0.12 Mg ha−1 yr−1) under 7 year study period. It is previously reported that ICL system contains labile organic matter pools10,38, subsequently showing higher C stocks and greater root densities near soil-surface, which promotes aggregate-associated C stabilization18,39,40, higher infiltration rates, thus providing likely benefits to soil function linked to erosion control and soil water relations41.Soil microbial community compositionTotal bacterial biomass, AM fungi, and PLFA were enhanced under NE, which can be result of accumulation of organic residues and higher pasture root mass7,32, pasture being grazed can promote exudation of organic compounds by roots, serving as energy sources for microorganisms. The consistent increase in microbial population under NE can also be result of increased SOC and N, however, the same does not hold true for ICL system, where despite observing greater SOC and N, a significant decrease in the microbial population at site 2 was noticed. The enhanced total PLFA under NE system at site 2 is due to concomitant increase in AM fungi, gram (−), fungal/bacterial ratio, and total bacterial biomass compared to ICL. The fungal to bacterial ratio was reduced under ICL compared to NE at sites 1 and 2, pertaining to relatively low abundance of the fungal fatty acid 18:2ɷ6 in grazed system as compared to unmanaged grassland. This finding corroborates the notion that livestock-grazing systems contain bacterial-based decomposition channels and are mostly dominated by gram (+) bacteria and that the fungal population is comparatively more important in decomposer food-webs of native grasslands. These results coincide with previous studies42,43. Moreover, the increase in fungal to bacterial ratios under NE system in contrast to ICL at sites 1 and 2 can relate to modifications in soil health with C sequestration, as fungal populations incline towards higher C assimilation proficiencies and greater storage of metabolized C than bacterial populations9,44. The grazing intensity also plays a significant role in bacterial and fungal presence. It is previously reported that high grazing intensity had greater bacterial PLFA concentration than the low grazing counterparts in grassland systems45. It is considered that under heavily grazed sites in grasslands, bacteria-based energy channels of decomposition dominate other microbial communities, while fungi can successfully enable decomposition in both slightly grazed and non-grazed systems43. Grazed pasture mix at short-term study site 4 showed 12–21% higher total PLFA than the oats, oats with CC, oats with CC + grazing systems. It is also possible that this increased total PLFA at site 4 under grazed pasture mix contributed to enhancing the 1–2 and 2–4 mm sized aggregates compared to other treatments. It indicated that though physicochemical properties can take longer ( > 8–15 years) in significantly responding to changes in management systems, soil microbial community and structure may show a rapid response (~ 3 years), thus it can be used as an early indicator while assessing the variations in soil health18,46.Overall, NE exemplified the undisturbed grazed mixture with a greater microbial population at sites 1, 2, and 3, when compared to other agricultural systems. Our findings coincide with previous studies where pasture systems performed better than the agricultural soil, in terms of, showing greater microbial biomass and fatty acid signatures related to bacterial and fungal populations, which is mostly attributed to greater surface coverage and absence of tillage practices in pasture systems9,47,48. Lower soil microbial communities under ICL system than native Cerrado pasture have been found previously because of reduced soil porosity and macropore continuity resulting in restricted gas diffusion and water movement18.Although the AM fungi abundance was not significant for sites 3 and 4, and significantly lower for ICL system than NE at sites 1 and 2, it should be taken into consideration that FAME analysis cannot reflect species-level changes for fungi and/or bacteria and the variations in microbial community structure for ICL system can be due to changes in abundance and distribution among microbial groups. For example, in a previous study9, while increased bacterial population was observed for continuous cotton compared to the ICL system, however, pyrosequencing for bacterial diversity assessment demonstrated disparities between both systems, where greater Proteobacteria was seen under ICL system than continuous cotton. Numerous factors such as degree of disturbance, pH level, bulk density, porosity, soil water content, C and N distribution, and residue positioning regulate the amount of bacterial and fungal biomass in agroecosystems18,49. Arbuscular mycorrhizal fungi are responsible for formation of macroaggregates ( > 0.25 mm) by producing a glycoprotein called glomalin, which is present abundantly in natural and agricultural systems. However, increased grazing intensity, use of excess fertilizers and fungicides can directly or indirectly reduce mycorrhizal population by influencing soil organisms accountable for converting soil organic matter into plant nutrients38. Animals may also cause moderate soil compaction affecting the fungal biodiversity and soil pore space6,38.Relationship among measured soil propertiesBased on PCA, it is derived that integrated crop–livestock and natural ecosystem of native grassland can provide substrate for the microbial composition and enhance aggregate-associated C and N fractions. A positive correlation between SOC and microbial communities suggested the inclination of microbes to affect SOC and N turnover and vice-versa through interaction with crop–livestock grazing, vegetation, and soil properties. Fungi exhibited insignificant responses to changes in soil pH and bulk density than bacteria because chitinous cell walls make fungi more resistant and resilient to variations in soil conditions50,51. A reduction in gram (−) bacteria may indicate the presence of stressed soil conditions due to pH and increased bulk density, which has previously been observed in other studies52,53. A significant negative correlation between bulk density and SOC, N, gram (−), actinomycetes, total bacteria, and total PLFA indicated that the microorganisms influenced the soil compaction and related SOC and N. Moreover, positive correlation between SOC and microbial composition suggested that microbes can influence C sequestration in the soil via a shift in community structure. Microbial composition is influenced by soil C, whereas N is a critical biogenic element that improves microbial growth and their ability to utilize soil C54. More