Philippa Kaur

1.
Hutchinson, G. E. Homage to Santa Rosalia or why are there so many kinds of animals?. Am. Nat. 93, 145–159 (1959).
Article  Google Scholar 
2.
Volterra, V. Variations and fluctuations of the number of individuals in animal species living together. ICES J. Mar. Sci. 3, 3–51 (1928).
Article  Google Scholar 

3.
MacArthur, R. & Levins, R. Competition, habitat selection, and character displacement in a patchy environment. Proc. Natl. Acad. Sci. USA. 51, 1207–1210 (1964).
ADS  CAS  PubMed  Article  PubMed Central  Google Scholar 

4.
Levin, S. A. Community equilibria and stability, and an extension of the competitive exclusion principle. Am. Nat. 104, 413–423 (1970).
Article  Google Scholar 

5.
Chesson, P. Mechanisms of maintenance of species diversity. Annu. Rev. Ecol. Syst. 31, 343–366 (2000).
Article  Google Scholar 

6.
Chase, J. M. et al. The interaction between predation and competition: a review and synthesis. Ecol. Lett. 5, 302–315 (2002).
Article  Google Scholar 

7.
Amarasekare, P. Competitive coexistence in spatially structured environments: a synthesis. Ecol. Lett. 6, 1109–1122 (2003).
Article  Google Scholar 

8.
Hutchinson, G. E. The paradox of the plankton. Am. Nat. 95, 137–145 (1961).
Article  Google Scholar 

9.
Connell, J. H. Diversity in tropical rain forests and coral reefs. Science 199, 1302–1310 (1978).
ADS  CAS  PubMed  Article  PubMed Central  Google Scholar 

10.
Armstrong, R. A. & McGehee, R. Competitive exclusion. Am. Nat. 115, 151–170 (1980).
MathSciNet  Article  Google Scholar 

11.
Huisman, J. & Weissing, F. J. Biodiversity of plankton by species oscillations and chaos. Nature 402, 407–410 (1999).
ADS  Article  Google Scholar 

12.
Abrams, P. A. & Holt, R. D. The impact of consumer–resource cycles on the coexistence of competing consumers. Theor. Popul. Biol. 62, 281–295 (2002).
PubMed  MATH  Article  Google Scholar 

13.
Schwartz, M. D. et al. Phenology: An Integrative Environmental Science (Kluwer Academic Publishers, New York, 2003).
Google Scholar 

14.
McMeans, B. C., McCann, K. S., Humphries, M., Rooney, N. & Fisk, A. T. Food web structure in temporally-forced ecosystems. Trends Ecol. Evol. 30, 662–672 (2015).
PubMed  Article  Google Scholar 

15.
White, E. R. & Hastings, A. Seasonality in Ecology: Progress and Prospects in Theory (Springer, New York, 2018).
Google Scholar 

16.
Rudolf, V. H. W. The role of seasonal timing and phenological shifts for species coexistence. Ecol. Lett. 22, 1324–1338 (2019).
PubMed  Google Scholar 

17.
Stewart, F. M. & Levin, B. R. Partitioning of resources and the outcome of interspecific competition: a model and some general considerations. Am. Nat. 107, 171–198 (1973).
Article  Google Scholar 

18.
Abrams, P. Variability in resource consumption rates and the coexistence of competing species. Theor. Popul. Biol. 25, 106–124 (1984).
MATH  Article  Google Scholar 

19.
Cushing, J. M. Periodic two-predator, one-prey interactions and the time sharing of a resource niche. SIAM J. Appl. Math. 44, 392–410 (1984).
MathSciNet  MATH  Article  Google Scholar 

20.
Grover, J. P. Resource competition in a variable environment: phytoplankton growing according to Monod’s model. Am. Nat. 136, 771–789 (1990).
Article  Google Scholar 

21.
Loreau, M. Time scale of resource dynamics and coexistence through time partitioning. Theor. Popul. Biol. 41, 401–412 (1992).
MATH  Article  Google Scholar 

22.
Namba, T. & Takahashi, S. Competitive coexistence in a seasonally fluctuating environment II. Multiple stable states and invasion success. Theor. Popul. Biol. 44, 374–402 (1993).
MathSciNet  MATH  Article  Google Scholar 

23.
Chesson, P. Multispecies competition in variable environments. Theor. Popul. Biol. 45, 227–276 (1994).
MATH  Article  Google Scholar 

24.
Abrams, P. A. When does periodic variation in resource growth allow robust coexistence of competing consumer species?. Ecology 85, 372–382 (2004).
Article  Google Scholar 

25.
Gravel, D., Guichard, F. & Hochberg, M. E. Species coexistence in a variable world. Ecol. Lett. 14, 828–839 (2011).
PubMed  Article  PubMed Central  Google Scholar 

26.
Sakavara, A., Tsirtsis, G., Roelke, D. L., Mancy, R. & Spatharis, S. Lumpy species coexistence arises robustly in fluctuating resource environments. Proc. Natl. Acad. Sci. USA. 115, 738–743 (2018).
CAS  PubMed  Article  PubMed Central  Google Scholar 

27.
Dunlap, J. C., Loros, J. J. & DeCoursey, P. J. Chronobiology: Biological Timekeeping (Sinauer Associates, London, 2004).
Google Scholar 

28.
Kronfeld-Schor, N. & Dayan, T. Partitioning of time as an ecological resource. Annu. Rev. Ecol. Evol. Syst. 34, 153–181 (2003).
Article  Google Scholar 

29.
Kronfeld-Schor, N. et al. Chronobiology by moonlight. Proc. R. Soc. Lond. B 280, 20123088 (2013).
Google Scholar 

30.
Welch, K. D. & Harwood, J. D. Temporal dynamics of natural enemy-pest interactions in a changing environment. Biol. Control 75, 18–27 (2014).
Article  Google Scholar 

31.
Raible, F., Takekata, H. & Tessmar-Raible, K. An overview of monthly rhythms and clocks. Front. Neurol. 8, 189 (2017).
PubMed  PubMed Central  Article  Google Scholar 

32.
Körtner, G. & Geiser, F. The temporal organization of daily torpor and hibernation: circadian and circannual rhythms. Chronobiol. Int. 17, 103–128 (2000).
PubMed  Article  Google Scholar 

33.
Holt, R. D. & Polis, G. A. A theoretical framework for intraguild predation. Am. Nat. 149, 745–764 (1997).
Article  Google Scholar 

34.
Holt, R. D. Predation, apparent competition, and the structure of prey communities. Theor. Popul. Biol. 12, 197–229 (1977).
MathSciNet  CAS  PubMed  Article  Google Scholar 

35.
Connell, J. H. Some mechanisms producing structure in natural communities: a model and evidence from field experiments. Ecol. Evol. Commun. 1, 460–490 (1975).
Google Scholar 

36.
Cozzi, G. et al. Fear of the dark or dinner by moonlight? Reduced temporal partitioning among africa’s large carnivores. Ecology 93, 2590–2599 (2012).
PubMed  Article  Google Scholar 

37.
Campera, M. et al. Temporal niche separation between the two ecologically similar nocturnal Primates Avahi meridionalis and Lepilemur fleuretae. Behav. Ecol. Sociobiol. 73, 1–10 (2019).
Article  Google Scholar 

38.
Leonard, J. P., Tewes, M. E., Lombardi, J. V., Wester, D. W. & Campbell, T. A. Effects of sun angle, lunar illumination, and diurnal temperature on temporal movement rates of sympatric ocelots and bobcats in South Texas. PLoS ONE 15, e0231732 (2020).
CAS  PubMed  PubMed Central  Article  Google Scholar 

39.
Shimadzu, H., Dornelas, M., Henderson, P. A. & Magurran, A. E. Diversity is maintained by seasonal variation in species abundance. BMC Biol. 11, 98 (2013).
PubMed  PubMed Central  Article  Google Scholar 

40.
Gaston, K. J., Bennie, J., Davies, T. W. & Hopkins, J. The ecological impacts of nighttime light pollution: a mechanistic appraisal. Biol. Rev. 88, 912–927 (2013).
PubMed  Article  PubMed Central  Google Scholar 

41.
Lovegrove, B. G. et al. Are tropical small mammals physiologically vulnerable to Arrhenius effects and climate change?. Physiol. Biochem. Zool. 87, 30–45 (2014).
PubMed  Article  PubMed Central  Google Scholar 

42.
Yerushalmi, S. & Green, R. M. Evidence for the adaptive significance of circadian rhythms. Ecol. Lett. 12, 970–981 (2009).
PubMed  Article  PubMed Central  Google Scholar 

43.
Bradshaw, W. E. & Holzapfel, C. M. Genetic response to rapid climate change: it’s seasonal timing that matters. Mol. Ecol. 17, 157–166 (2008).
CAS  PubMed  Article  PubMed Central  Google Scholar 

44.
Sauve, D., Divoky, G. & Friesen, V. L. Phenotypic plasticity or evolutionary change? An examination of the phenological response of an arctic seabird to climate change. Funct. Ecol. 33, 2180–2190 (2019).
Article  Google Scholar 

45.
Abbey-Lee, R. N. & Dingemanse, N. J. Adaptive individual variation in phenological responses to perceived predation levels. Nat. Commun. 10, 1601 (2019).
ADS  CAS  PubMed  PubMed Central  Article  Google Scholar  More

1.
Smith, S. E. & Read, D. J. Mycorrhizal Symbiosis 3rd edn. (Academic Press, London, 2008).
Google Scholar 
2.
Gianinazzi, S. et al. Agroecology: the key role of arbuscularmycorrhizas in ecosystem services. Mycorrhiza 20, 519–530 (2010).
Article  Google Scholar 

3.
Lenoir, I., Fontaine, J. & Sahraoui, A. L. H. Arbuscularmycorrhizal fungal responses to abiotic stresses: a review. Phytochem 123, 4–15 (2016).
CAS  Article  Google Scholar 

4.
Song, Y., Chen, D., Lu, K., Sun, Z. & Zeng, R. Enhanced tomato disease resistance primed by arbuscularmycorrhizal fungus. Front. Plant Sci. 6, 786 (2015).
PubMed  PubMed Central  Google Scholar 

5.
Van Geel, M. et al. Abiotic rather than biotic filtering shapes the arbuscularmycorrhizal fungal communities of European seminatural grasslands. New Phytol. 220, 1262–1272 (2018).
Article  Google Scholar 

6.
Varma, A., Prasad, R. & Tuteja, N. Mycorrhiza—Nutrient Uptake (Biocontrol, Ecorestoration Fourth Edition, Springer, 2017).
Google Scholar 

7.
Yu, L., Nicolaisen, J., Larsen, J. & Ravnskov, S. Molecular characterization of root-associated fungal communities in relation to health status of Pisum sativum using barcoded pyrosequencing. Plant Soil 357, 395–405 (2012).
CAS  Article  Google Scholar 

8.
Corredor, A. H., Van Rees, K. & Vujanovic, V. Host genotype and health status influence on the composition of the arbuscularmycorrhizal fungi in Salix bioenergy plantations. For. Ecol. Manag. 314, 112–119 (2014).
Article  Google Scholar 

9.
Martinez, N. & Johnson, N. C. Agricultural management influences propagule densities and functioning of arbuscularmycorrhizas in low- and high-input agroecosystems in arid environments. Appl. Soil Ecol. 46, 300–306 (2010).
Article  Google Scholar 

10.
Hontoria, C., García-González, I., Quemada, M., Roldánd, A. & Alguacil, M. M. The cover crop determines the AMF community composition in soil and in roots of maize after a ten-year continuous crop rotation. Sci. Total Environ. 660, 913–922 (2019).
ADS  CAS  Article  Google Scholar 

11.
Lumini, E., Vallino, M., Alguacil, M. M., Romani, M. & Bianciotto, V. Different farming and water regimes in Italian rice fields affect arbuscularmycorrhizal fungal soil communities. Ecol. Appl. 21, 1696–1707 (2011).
Article  Google Scholar 

12.
Manoharan, L., Rosenstock, N. P., Williams, A. & Hedlund, K. Agricultural management practices influence AMF diversity and community composition with cascading effects on plant productivity. Appl. Soil Ecol. 115, 53–59 (2017).
Article  Google Scholar 

13.
Dai, M., Bainard, L. D., Hamel, C., Gan, Y. & Lynch, D. Impact of land use on arbuscularmycorrhizal fungal communities in rural Canada. Appl. Environ. Microbiol. 79, 6719–6729 (2013).
CAS  Article  Google Scholar 

14.
Aghili, F. et al. Wheat plants invest more in mycorrhizae and receive more benefits from them under adverse than favorable soil conditions. Appl. Soil Ecol. 84, 93–111 (2014).
Article  Google Scholar 

15.
Gaudin, J., Semetey, O., Foissac, X. & Eveillard, S. Phytoplasmatiter in diseased lavender is not correlated to lavender tolerance to stolburphytoplasma. Bull. Insectol. 64(Supplement), S179–S180 (2011).
Google Scholar 

16.
Kamińska, M., Klamkowski, K., Berniak, H. & Treder, W. Effect of arbuscularmycorrhizal fungi inoculation on aster yellows phytoplasma-infected tobacco plants. Sci. Hortic. 125, 500–503 (2010).
Article  Google Scholar 

17.
Batlle, A. et al. Tolerance increase to Candidatus phytoplasma prunorum in mycorrhizal plums fruit trees. Bull. Insectol. 64, 125–126 (2011).
Google Scholar 

18.
D’ameli, R. et al. Increased plant tolerance against chrysanthemum yellows phytoplasma (Candidatus Phytoplasma asteris) following double inoculation with Glomusmosseae BEG12 and Pseudomonas putida S1Pf1Rif. Plant. Pathol. 60, 1014–1022 (2011).
Article  Google Scholar 

19.
Fiorilli, V. et al. Omics approaches revealed how arbuscularmycorrhizal symbiosis enhances yield and resistance to leaf pathogen in wheat. Sci. Rep. 8, 9625 (2018).
ADS  Article  Google Scholar 

20.
Bødker, L., Kjøller, R., Kristensen, K. & Rosendahl, S. Interactions between indigenous arbuscularmycorrhizal fungi and Aphanomyces euteiches in field-grown pea. Mycorrhiza 12, 7–12 (2002).
Article  Google Scholar 

21.
Al-Askar, A. A. & Rashad, Y. M. Arbuscularmycorrhizal fungi: a biocontrol agent against common bean Fusarium root rot disease. Plant Pathol. J. 9, 31–38 (2010).
Article  Google Scholar 

22.
Hugoni, M., Luis, P., Guyonnet, J. & Haichar, F. Z. Plant host habitat and root exudates shape fungal diversity. Mycorrhiza 28, 451–463 (2018).
Article  Google Scholar 

23.
Bertaccini, A. & Duduk, B. Phytoplasma and phytoplasma diseases: A review of recent research. Phytopathol. Mediterr. 48, 355–378 (2009).
CAS  Google Scholar 

24.
Stierlin, E., Nicolè, F., Costes, T., Fernandez, X. & Michel, T. Metabolomic study of volatile compounds emitted by lavender grown under open-field conditions: a potential approach to investigate the yellow decline disease. Metabolomics 16, 31 (2020).
CAS  Article  Google Scholar 

25.
Lopez-Garcia, A. et al. Plant traits determine the phylogenetic structure of arbuscularmycorrhizal fungal communities. Mol. Ecol. 26, 6948–6959 (2017).
Article  Google Scholar 

26.
Alguacil, M. M., Díaz, G., Torres, P., Rodríguez-Caballero, G. & Roldan, A. Host identity and functional traits determine the community composition of the arbuscularmycorrhizal fungi in facultative epiphytic plant species. Fungal Ecol. 39, 307–315 (2019).
Article  Google Scholar 

27.
Neuenkamp, L. et al. The role of plant mycorrhizal type and status in modulating the relationship between plant and arbuscularmycorrhizal fungal communities. New Phytol. 220, 1236–1247 (2018).
CAS  Article  Google Scholar 

28.
Alguacil, M. M., Torrecillas, E., García-Orenes, F. C. & Roldán, A. Changes in the composition and diversity of AMF communities mediated by management practices in a Mediterranean soil are related with increases in soil biological activity. Soil Biol. Biochem. 76, 34–44 (2014).
CAS  Article  Google Scholar 

29.
Giri, B. & Mukerji, K. G. Mycorrhizal inoculant alleviates salt stress in Sesbania aegyptiaca and Sesbania grandiflora under field conditions: evidence for reduced sodium and improved magnesium uptake. Mycorrhiza 14, 307–312 (2004).
Article  Google Scholar 

30.
Phillips, J. M. & Hayman, D. S. Improved procedure for clearing roots and staining parasitic and vesicular–arbuscularmycorrhizal fungi for rapid assessment of infection. Trans. Br. Mycol. Soc. 55, 158–163 (1970).
Article  Google Scholar 

31.
Trouvelot, A., Kough, J. L. & Gianinazzi-Pearson, V. Mesure du taux de mycorhization VA d’un système radiculaire. Recherche de méthodes ayant une signification fonctionnelle. In Physiological and genetical aspects of mycorrhizae (eds Gianinazzi-Pearson, V. & Gianinazzi, S.) 217–221 (INRA Press, Paris, 1986).
Google Scholar 

32.
Gollotte, A., van Tuinen, D. & Atkinson, D. Diversity of arbuscularmycorrhizalfungicolonisingroots of the grassspeciesAgrostis capillaris and Lolium perenne in a fieldexperiment. Mycorrhiza 14, 111–117 (2004).
Article  Google Scholar 

33.
Binet, M. N. et al. Responses of above- and below-ground fungal symbionts to cessation of mowing in subalpine grassland. Fungal Ecol. 25, 14–21 (2017).
Article  Google Scholar 

34.
Mouhamadou, B. et al. Effects of two grass species on the composition of soil fungal communities. Biol. Fertil. Soils 49, 1131–1139 (2013).
Article  Google Scholar 

35.
Boyer, F. et al. Obitools: a unix- inspired software package for DNA metabarcoding. Mol. Ecol. Resour. 16, 176–182 (2016).
CAS  Article  Google Scholar 

36.
Lentendu, G. et al. Assessment of soil fungal diversity in different alpine tundra habitats by means of pyrosequencing. Fungal Div. 49, 113–123 (2011).
Article  Google Scholar 

37.
van Dongen, S. Graph clustering by flow simulation. Ph.D. thesis, University of Utrecht (2000).

38.
Thompson, L. A. S-PLUS (and R) manual to accompany Agresti’s Categorical Data Analysis (2002), 2nd ed (2009).

39.
Oksanen, J., Kindt, R., Legendre, P., O’Hara, B. & Gavin, L. vegan: Community Ecology Package. R package version 1.15–4 (2009).

40.
Mouhamadou, B. et al. Molecular screening of xerophilic Aspergillus strains producing mycophenolic acid. Fungal Biol. 121, 103–111 (2017).
CAS  Article  Google Scholar  More

1.
Xu, X. et al. Interannual variability in responses of belowground net primary productivity (NPP) and NPP partitioning to long-term warming and clipping in a tallgrass prairie. Global Change Biol. 18(5), 1648–1656 (2012).
ADS  Article  Google Scholar 
2.
Hui, D. & Jackson, R. B. Geographical and interannual variability in biomass partitioning in grassland ecosystems: a synthesis of field data. New Phytol. 169(1), 85–93 (2006).
CAS  Article  Google Scholar 

3.
Zhang, X. et al. Impact of prolonged drought on rainfall use efficiency using MODIS data across China in the early 21st century. Remote Sens. Environ. 150, 188–197 (2014).
ADS  Article  Google Scholar 

4.
Jiang, Y. et al. Effects of community structure on precipitation-use efficiency of grasslands in Northern Tibet. J. Veg Sci. 28, 281–290 (2017).
Article  Google Scholar 

5.
Roupsard, O. et al. Scaling-up productivity (NPP) using light or water use efficiencies (LUE, WUE) from a two-layer tropical plantation. Agrofor. Syst. 76(2), 409–422 (2009).
Article  Google Scholar 

6.
Prince, S. D., De Colstoun, E. B. & Kravitz, L. L. Evidence from rain-use efficiencies does not indicate extensive Sahelian desertification. Global Change Biol. 4(4), 359–374 (1998).
ADS  Article  Google Scholar 

7.
Fensholt, R. & Rasmussen, K. Analysis of trends in the Sahelian “rain-use efficiency” using GIMMS NDVI, RFE and GPCP rainfall data. Remote Sens. Environ. 115(2), 438–451 (2011).
ADS  Article  Google Scholar 

8.
Ye, H., Wang, J., Huang, M. & Qi, S. Spatial pattern of vegetation precipitation use efficiency and its response to precipitation and temperature on the Qinghai-Xizang Plateau of China. Chin. J. Plant. Ecol. 36(12), 1237–1247 (2012).
Article  Google Scholar 

9.
Bai, Y. et al. Primary production and rain use efficiency across a precipitation gradient on the Mongolia plateau. Ecology 89(8), 2140–2153 (2008).
Article  Google Scholar 

10.
Paruelo, J. M., Lauenroth, W. K., Burke, I. C. & Sala, O. E. Grassland precipitation-use efficiency varies across a resource gradient. Ecosystems 2(1), 64–68 (1999).
Article  Google Scholar 

11.
Yang, Y., Fang, J., Fay, P. A., Bell, J. E. & Ji, C. Rain use efficiency across a precipitation gradient on the Tibetan Plateau. Geophys. Res. Lett. 37, L15702 (2010).
ADS  Google Scholar 

12.
Li, H. X., Liu, G. H. & Fu, B. J. Spatial variations of rain-use efficiency along a climate gradient on the Tibetan Plateau: a satellite-based analysis. Int. J. Remote Sens. 34(21), 7487–7503 (2013).
ADS  Article  Google Scholar 

13.
Huxman, T. E. et al. Convergence across biomes to a common rain-use efficiency. Nature 429(6992), 651–654 (2004).
ADS  CAS  Article  Google Scholar 

14.
Hu, Z. et al. Precipitation-use efficiency along a 4500-km grassland transect. Glob. Ecol. Biogeogr. 19(6), 842–851 (2010).
Article  Google Scholar 

15.
Lauenroth, W. K., Burke, I. C. & Paruelo, J. M. Patterns of production and precipitation-use efficiency of winter wheat and native grasslands in the central Great Plains of the United States. Ecosystems 3(4), 344–351 (2000).
Article  Google Scholar 

16.
Hooper, D. U. & Johnson, L. Nitrogen limitation in dryland ecosystems: Responses to geographical and temporal variation in precipitation. Biogeochemistry 46(1–3), 247–293 (1999).
CAS  Google Scholar 

17.
Xu, X., Sherry, R. A., Niu, S., Li, D. & Luo, Y. Net primary productivity and rain-use efficiency as affected by warming, altered precipitation, and clipping in a mixed-grass prairie. Global Change Biol. 19(9), 2753–2764 (2013).
ADS  Article  Google Scholar 

18.
De Boeck, H. J. et al. How do climate warming and plant species richness affect water use in experimental grasslands?. Plant Soil 288(1–2), 249–261 (2006).
ADS  Article  Google Scholar 

19.
Campos, G. E. P. et al. Ecosystem resilience despite large-scale altered hydroclimatic conditions. Nature 494(7437), 349–352 (2013).
ADS  Article  Google Scholar 

20.
Qiu, J., Zhang, H. & Shen, W. Spatial characteristics of precipitation use efficiency on the Qinghai-Tibet Plateau From 1982 to 2007. J. Fudan. Univ. Nat. Sci. 53(1), 126–133 (2014).
Google Scholar 

21.
Wang, Q. W., Yu, D. P., Dai, L. M., Zhou, L. & Zhou, W. M. Research progress in water use efficiency of plants under global climate change. Chin. J. Appl. Ecol. 21(12), 3255–3265 (2000).
Google Scholar 

22.
Chen, S. P., Bai, Y. F., Zhang, L. X. & Han, X. G. Comparing physiological responses of two dominant grass species to nitrogen addition in Xilin River Basin of China. Environ. Exp. Bot. 53(1), 65–75 (2005).
Article  Google Scholar 

23.
Qiu, J. The third pole. Nature 454(7203), 393–396 (2008).
CAS  Article  Google Scholar 

24.
Chen, B. X. et al. The impact of climate change and anthropogenic activities on alpine grassland over the Qinghai-Tibet Plateau. Agric. For. Meteorol. 189, 11–18 (2014).
ADS  Article  Google Scholar 

25.
Jiang, Y. B. et al. Effects of community structure on precipitation-use efficiency of grasslands in northern Tibet. J. Veg. Sci. 28, 281–290 (2017).
Article  Google Scholar 

26.
Gao, Q. Z. et al. Effects of topography and human activity on the net primary productivity (NPP) of alpine grassland in northern Tibet from 1981 to 2004. Int. J. Remote Sens. 34(6), 2057–2069 (2013).
ADS  Article  Google Scholar 

27.
Zhang, J. H., Yao, F. M., Zheng, L. G. & Yang, L. M. Evaluation of grassland dynamics in the Northern-Tibet Plateau of China using remote sensing and climate data. Sensors 7(12), 3312–3328 (2007).
Article  Google Scholar 

28.
Li, Z., Huffman, T., McConkey, B. & Townley-Smith, L. Monitoring and modeling spatial and temporal patterns of grassland dynamics using time-series MODIS NDVI with climate and stocking data. Remote Sens. Environ. 138, 232–244 (2013).
ADS  Article  Google Scholar 

29.
Zhang, X. K., Lu, X. Y. & Wang, X. D. Spatial-temporal NDVI variation of different alpine grassland classes and groups in Northern Tibet from 2000 to 2013. Mt. Res. Dev. 35(3), 254–263 (2015).
Article  Google Scholar 

30.
Yu, D. Y., Shi, P. J., Shao, H. B., Zhu, W. Q. & Pan, Y. H. Modelling net primary productivity of terrestrial ecosystems in East Asia based on an improved CASA ecosystem model. Int. J. Remote Sens. 30(18), 4851–4866 (2009).
ADS  Article  Google Scholar 

31.
Gao, Q. Z. et al. Dynamics of alpine grassland NPP and its response to climate change in Northern Tibet. Clim. Change 97(3–4), 515–528 (2009).
ADS  CAS  Article  Google Scholar 

32.
Zhu, W. Q., Pan, Y. Z., He, H., Yu, D. Y. & Hu, H. B. Simulation of maximum light use efficiency for some typical vegetation types in China. Chin. Sci. Bull. 51(4), 457–463 (2006).
CAS  Article  Google Scholar 

33.
Zhao, G. S. et al. Spatial-temporal variation of ANPP and rain-use efficiency along a precipitation gradient on Changtang Plateau, Tibet. Remote Sens. 11, 325 (2019).
ADS  Article  Google Scholar 

34.
Sun, J. & Du, W. Effects of precipitation and temperature on net primary productivity and precipitation use efficiency across China’s grasslands. GISci. Remote Sens. 54(6), 881–897 (2017).
Article  Google Scholar 

35.
Chen, Z. Q., Shao, Q. Q., Liu, J. Y. & Wang, J. B. Analysis of net primary productivity of terrestrial vegetation on the Qinghai-Tibet Plateau, based on MODIS remote sensing data. Sci. China Earth Sci. 55(8), 1306–1312 (2012).
ADS  Article  Google Scholar 

36.
Piao, S. & Fang, J. Terrestrial net primary production and its spatio-temporal patterns in Qinghai-Xizang Plateau, China during 1982–1999. J. Nat. Resour. 03, 373–380 (2002).
Google Scholar 

37.
Gao, Q. Z., Wan, Y. F., Li, Y. E., Lin, E. D. & Yang, K. Grassland net primary production and its spatiotemporal distribution in Northern Tibet: a study with CASA model. Chin. J. Appl. Ecol. 11, 2526–2532 (2007).
Google Scholar 

38.
Zhou, C. P., Ouyang, H., Wang, Q. X., Watanabe, M. & Sun, Q. Q. Estimation net primary productivity in Tibetan Plateau. Acta Geogr. Sin. 01, 74–79 (2004).
Google Scholar 

39.
Yang, Y. H. et al. Storage, patterns and controls of soil organic carbon in the Tibetan grasslands. Global Change Biol. 14, 1592–1599 (2008).
ADS  Article  Google Scholar  More

1.
Spatafora, J. W. et al. A phylum-level phylogenetic classification of zygomycete fungi based on genome-scale data. Mycologia 108, 1028–1046 (2016).
CAS  PubMed  PubMed Central  Article  Google Scholar 
2.
Smith, S. E. & Read, D. J. Mycorrhizal Symbiosis (Academic Press, Amsterdam, 2008).
Google Scholar 

3.
van der Heijden, M. G. A., Martin, F. M., Selosse, M. A. & Sanders, I. R. Mycorrhizal ecology and evolution: the past, the present, and the future. New Phytol. 205, 1406–1423 (2015).
PubMed  Article  CAS  PubMed Central  Google Scholar 

4.
Lekberg, Y., Hammer, E. C. & Olsson, P. A. Plants as resource islands and storage units—adopting the mycocentric view of arbuscular mycorrhizal networks. FEMS Microbiol. Ecol. 74, 336–345 (2010).
CAS  PubMed  Article  PubMed Central  Google Scholar 

5.
Allen, M. F. Mycorrhizal fungi: highways for water and nutrients in arid soils. Vadose Zone J. 6, 291–297 (2007).
Article  Google Scholar 

6.
Newsham, K. K., Fitter, A. H. & Watkinson, A. R. Arbuscular mycorrhiza protect an annual grass from root pathogenic fungi in the field. J. Ecol. 83, 991–1000 (1995).
Article  Google Scholar 

7.
Vigo, C., Norman, J. R. & Hooker, J. E. Biocontrol of the pathogen Phytophthora parasitica by arbuscular mycorrhizal fungi is a consequence of effects on infection loci. Plant Pathol. 49, 509–514 (2000).
Article  Google Scholar 

8.
Aroca, R., Porcel, R. & Ruiz-Lozano, J. M. How does arbuscular mycorrhizal symbiosis regulate root hydraulic properties and plasma membrane aquaporins in Phaseolus vulgaris under drought, cold or salinity stresses?. New Phytol. 173(4), 808–816 (2007).
CAS  PubMed  Article  PubMed Central  Google Scholar 

9.
Augé, R. M., Toler, H. D. & Saxton, A. M. Arbuscular mycorrhizal symbiosis and osmotic adjustment in response to NaCl stress: a meta-analysis. Front Plant Sci. 5, ARTN 562. https://doi.org/10.3389/fpls.2014.00562 (2014).

10.
Augé, R. M., Toler, H. D. & Saxton, A. M. Arbuscular mycorrhizal symbiosis alters stomatal conductance of host plants more under drought than under amply watered conditions: a meta-analysis. Mycorrhiza 25(1), 13–24 (2015).
PubMed  Article  PubMed Central  Google Scholar 

11.
Pfeffer, P. E., Douds, D. D., Becard, G. & Shachar-Hill, Y. Carbon uptake and the metabolism and transport of lipids in an arbuscular mycorrhiza. Plant Physiol. 120(2), 587–598 (1999).
CAS  PubMed  PubMed Central  Article  Google Scholar 

12.
Bago, B., Pfeffer, P. E. & Shachar-Hill, Y. Carbon metabolism and transport in arbuscular mycorrhizas. Plant Physiol. 124(3), 949–958 (2000).
CAS  PubMed  PubMed Central  Article  Google Scholar 

13.
Horton, T. R. Mycorrhizal networks (Springer, Dordrecht, 2015).
Google Scholar 

14.
Walder, F. & van der Heijden, M. G. A. Regulation of resource exchange in the arbuscular mycorrhizal symbiosis. Nat. Plants 1(11), 7 (2015).
Article  CAS  Google Scholar 

15.
van der Heijden, M. G. A. et al. Mycorrhizal fungal diversity determines plant biodiversity, ecosystem variability and productivity. Nature 396(6706), 69–72 (1998).
ADS  Article  CAS  Google Scholar 

16.
Wilson, G. W. T., Hartnett, D. C. & Rice, C. W. Mycorrhizal-mediated phosphorus transfer between the tallgrass prairie plants Sorghastrum nutans and Artemisia ludoviciana. Funct. Ecol. 20, 427–435 (2006).
Article  Google Scholar 

17.
Bever, J. D. et al. Rooting theories of plant community ecology in microbial interactions. Trends Ecol. Evol. 25(8), 468–478 (2010).
PubMed  PubMed Central  Article  Google Scholar 

18.
Walder, F. et al. Mycorrhizal networks: common goods of plants shared under unequal terms of trade. Plant Physiol. 159, 789–797 (2012).
CAS  PubMed  PubMed Central  Article  Google Scholar 

19.
Weremijewicz, J., Sternberg, L. & Janos, D. P. Common mycorrhizal networks amplify competition by preferential mineral nutrient allocation to large host plants. New Phytol. 212(2), 461–471 (2016).
CAS  PubMed  Article  PubMed Central  Google Scholar 

20.
Řezáčová, V. et al. Little cross-feeding of the mycorrhizal networks shared between C3-Panicum bisulcatum and C4-Panicum maximum under different temperature regimes. Front. Plant Sci. 9, 16. https://doi.org/10.3389/fpls.2018.00449 (2018).
Article  Google Scholar 

21.
Deslippe, J. R. & Simard, S. W. Below-ground carbon transfer among Betula nana may increase with warming in Arctic tundra. New Phytol. 192, 689–698 (2011).
CAS  PubMed  Article  PubMed Central  Google Scholar 

22.
Bever, J. D., Richardson, S. C., Lawrence, B. M., Holmes, J. & Watson, M. Preferential allocation to beneficial symbiont with spatial structure maintains mycorrhizal mutualism. Ecol. Lett. 12(1), 13–21 (2009).
PubMed  Article  PubMed Central  Google Scholar 

23.
Lendenmann, M. et al. Symbiont identity matters: carbon and phosphorus fluxes between Medicago truncatula and different arbuscular mycorrhizal fungi. Mycorrhiza 21(8), 689–702 (2011).
CAS  PubMed  Article  PubMed Central  Google Scholar 

24.
Kiers, E. T. et al. Reciprocal rewards stabilize cooperation in the mycorrhizal symbiosis. Science 333(6044), 880–882 (2011).
ADS  CAS  PubMed  Article  PubMed Central  Google Scholar 

25.
Rillig, M. C. Arbuscular mycorrhizae and terrestrial ecosystem processes. Ecol. Lett. 7, 740–754 (2004).
Article  Google Scholar 

26.
Verbruggen, E. & Kiers, E. T. Evolutionary ecology of mycorrhizal functional diversity in agricultural systems. Evol Appl. 3(5–6), 547–560 (2010).
PubMed  PubMed Central  Article  Google Scholar 

27.
van Kleunen, M. et al. Global exchange and accumulation of non-native plants. Nature 525(7567), 100–103 (2015).
ADS  PubMed  Article  CAS  PubMed Central  Google Scholar 

28.
Pejchar, L. & Mooney, H. A. Invasive species, ecosystem services and human well-being. Trends Ecol. Evol. 24(9), 497–504 (2009).
PubMed  Article  PubMed Central  Google Scholar 

29.
Pyšek, P. et al. A global assessment of invasive plant impacts on resident species, communities and ecosystems: the interaction of impact measures, invading species’ traits and environment. Glob. Change Biol. 18(5), 1725–1737 (2012).
ADS  Article  Google Scholar 

30.
Blackburn, T. M. et al. A unified classification of alien species based on the magnitude of their environmental impacts. PLoS Biol. 12(5), ARTN e1001850. https://doi.org/10.1371/journal.pbio.1001850 (2014).

31.
Mitchell, C. E. et al. Biotic interactions and plant invasions. Ecol. Lett. 9(6), 726–740 (2006).
PubMed  Article  PubMed Central  Google Scholar 

32.
Catford, J. A., Jansson, R. & Nilsson, C. Reducing redundancy in invasion ecology by integrating hypotheses into a single theoretical framework. Divers. Distrib. 15(1), 22–40 (2009).
Article  Google Scholar 

33.
van der Putten, W. H. Impacts of soil microbial communities on exotic plant invasions. Trends Ecol. Evol. 25(9), 512–519 (2010).
PubMed  Article  Google Scholar 

34.
Keane, R. M. & Crawley, M. J. Exotic plant invasions and the enemy release hypothesis. Trends Ecol. Evol. 17(4), 164–170 (2002).
Article  Google Scholar 

35.
Pyšek, P. et al. Naturalization of central European plants in North America: species traits, habitats, propagule pressure, residence time. Ecology 96(3), 762–774 (2015).
PubMed  Article  Google Scholar 

36.
Davis, M. A., Grime, J. P. & Thompson, K. Fluctuating resources in plant communities: a generaltheory of invasibility. J. Ecol. 88, 528–534 (2000).
Article  Google Scholar 

37.
Callaway, R. M., Thelen, G. C., Rodriguez, A. & Holben, W. E. Soil biota and exotic plant invasion. Nature 427(6976), 731–733 (2004).
ADS  CAS  PubMed  Article  Google Scholar 

38.
Rudgers, J. A. & Orr, S. Non-native grass alters growth of native tree species via leaf and soil microbes. J. Ecol 97(2), 247–255 (2009).
Article  Google Scholar 

39.
Sun, Z. K. & He, W. M. Evidence for enhanced mutualism hypothesis: Solidago canadensis plants from regular soils perform better. PLoS ONE 5(11), 5. https://doi.org/10.1371/journal.pone.0015418 (2010).
CAS  Article  Google Scholar 

40.
Dickie, I. A. et al. The emerging science of linked plant-fungal invasions. New Phytol. 215(4), 1314–1332 (2017).
CAS  PubMed  Article  PubMed Central  Google Scholar 

41.
Cronk, Q. C. B. & Fuller, J. R. Plant Invaders: The Threat to Natural Ecosystems (Earthscan Publications, London, 2001).
Google Scholar 

42.
Richardson, D. M., Allsopp, N., D’Antonio, C. M., Milton, S. J. & Rejmanek, M. Plant invasions—the role of mutualisms. Biol. Rev. 75(1), 65–93 (2000).
CAS  PubMed  Article  PubMed Central  Google Scholar 

43.
Pringle, A. et al. Mycorrhizal symbioses and plant invasions. Ann Rev. Ecol. Evol. Syst. 40, 699–715 (2009).
Article  Google Scholar 

44.
Wilson, G. W. T., Hickman, K. R. & Williamson, M. M. Invasive warm-season grasses reduce mycorrhizal root colonization and biomass production of native prairie grasses. Mycorrhiza 22, 327–336 (2012).
PubMed  Article  PubMed Central  Google Scholar 

45.
Nunez, M. A. & Dickie, I. A. Invasive belowground mutualists of woody plants. Biol. Invasions 16, 645–661 (2014).
Article  Google Scholar 

46.
Bunn, R. A., Ramsey, P. W. & Lekberg, Y. Do native and invasive plants differ in their interactions with arbuscular mycorrhizal fungi? A meta-analysis. J. Ecol. 103, 1547–1556 (2015).
CAS  Article  Google Scholar 

47.
Gucwa-Przepiora, E., Chmura, D. & Sokolowska, K. AM and DSE colonization of invasive plants in urban habitat: a study of Upper Silesia (southern Poland). J. Plant Res. 129, 603–614 (2016).
PubMed  PubMed Central  Article  Google Scholar 

48.
Waller, L. P., Callaway, R. M., Klironomos, J. N., Ortega, Y. K. & Maron, J. L. Reduced mycorrhizal responsiveness leads to increased competitive tolerance in an invasive exotic plant. J. Ecol. 104, 1599–1607 (2016).
Article  Google Scholar 

49.
Menzel, A. et al. Mycorrhizal status helps explain invasion success of alien plant species. Ecology 98, 92–102 (2017).
PubMed  Article  PubMed Central  Google Scholar 

50.
Broadbent, A. A. D., Stevens, C. J., Ostle, N. J. & Orwin, K. H. Biogeographic differences in soil biota promote invasive grass response to nutrient addition relative to co-occurring species despite lack of belowground enemy release. Oecologia 186, 611–620 (2018).
ADS  PubMed  Article  PubMed Central  Google Scholar 

51.
Vogelsang, K. M. & Bever, J. D. Mycorrhizal densities decline in association with nonnative plants and contribute to plant invasion. Ecology 90, 399–407 (2009).
PubMed  Article  PubMed Central  Google Scholar 

52.
Reinhart, K. O. & Callaway, R. M. Soil biota and invasive plants. New Phytol. 170, 445–457 (2006).
PubMed  Article  PubMed Central  Google Scholar 

53.
Pakpour, S. & Klironomos, J. The invasive plant, Brassica nigra, degrades local mycorrhizas across a wide geographical landscape. R. Soc. Open Sci. 2, 4 (2015).
Article  Google Scholar 

54.
Shah, M. A., Reshi, Z. A. & Khasa, D. P. Arbuscular mycorrhizas: Drivers or passengers of alien plant invasion. Bot. Rev. 75, 397–417 (2009).
Article  Google Scholar 

55.
De Souza, T. A. F., Rodriguez-Echeverria, S., de Andrade, L. A. & Freitas, H. Could biological invasion by Cryptostegia madagascariensis alter the composition of the arbuscular mycorrhizal fungal community in semi-arid Brazil?. Acta Bot. Bras. 30, 93–101 (2016).
Article  Google Scholar 

56.
Awaydul, A. et al. Common mycorrhizal networks influence the distribution of mineral nutrients between an invasive plant, Solidago canadensis, and a native plant, Kummerowa striata. Mycorrhiza 29, 29–38 (2019).
CAS  PubMed  Article  PubMed Central  Google Scholar 

57.
Štajerová, K., Šmilauerová, M. & Šmilauer, P. Arbuscular mycorrhizal symbiosis of herbaceous invasive neophytes in the Czech Republic. Preslia 81, 341–355 (2009).
Google Scholar 

58.
Hempel, S. et al. Mycorrhizas in the Central European flora: relationships with plant life history traits and ecology. Ecology 94, 1389–1399 (2013).
PubMed  Article  PubMed Central  Google Scholar 

59.
Callaway, R. M., Newingham, B., Zabinski, C. A. & Mahall, B. E. Compensatory growth and competitive ability of an invasive weed are enhanced by soil fungi and native neighbours. Ecol. Lett. 4, 429–433 (2001).
Article  Google Scholar 

60.
Workman, R. E. & Cruzan, M. B. Common mycelial networks impact competition in an invasive grass. Am. J. Bot. 103, 1041–1049 (2016).
CAS  PubMed  Article  PubMed Central  Google Scholar 

61.
Zhang, Q. et al. Potential allelopathic effects of an invasive species Solidago canadensis on the mycorrhizae of native plant species. Allelopathy J. 20, 71–77 (2007).
ADS  CAS  Google Scholar 

62.
Callaway, R. M. et al. Novel weapons: Invasive plant suppresses fungal mutualists in America but not in its native Europe. Ecology 89, 1043–1055 (2008).
PubMed  Article  PubMed Central  Google Scholar 

63.
Sarma, K. K. V. Allelopathic potential of Echinops echinatus and Solanum surratense on seed germination of Argemone mexicana. Trop. Ecol. 15, 156–157 (1974).
Google Scholar 

64.
Smith, M. D., Hartnett, D. C. & Wilson, G. W. T. Interacting influence of mycorrhizal symbiosis and competition on plant diversity in tallgrass prairie. Oecologia 121, 574–582 (1999).
ADS  CAS  PubMed  Article  PubMed Central  Google Scholar 

65.
Bennett, J. A. et al. Plant-soil feedbacks and mycorrhizal type influence temperate forest population dynamics. Science 355, 181–184 (2017).
ADS  CAS  PubMed  Article  PubMed Central  Google Scholar 

66.
Liao, H. X. et al. Soil microbes regulate forest succession in a subtropical ecosystem in China: evidence from a mesocosm experiment. Plant Soil 430, 277–289 (2018).
CAS  Article  Google Scholar 

67.
Řezáčová, V. et al. Mycorrhizal symbiosis induces plant carbon reallocation differently in C3 and C4Panicum grasses. Plant Soil 425, 441–456 (2018).
Article  CAS  Google Scholar 

68.
Newman, E. I. A method of estimating total length of root in a sample. J. Appl. Ecol. 3, 139–145 (1966).
Article  Google Scholar 

69.
Bukovská, P., Gryndler, M., Gryndlerová, H., Püschel, D. & Jansa, J. Organic nitrogen-driven stimulation of arbuscular mycorrhizal fungal hyphae correlates with abundance of ammonia oxidizers. Front. Microbiol. 7, 711 (2016).
PubMed  PubMed Central  Article  Google Scholar 

70.
Hewitt, E. J. Sand and water culture methods used in the study of plant nutrition. CAB Tech. Commun. 22, 431–432 (1966).
Google Scholar 

71.
Řezáčová, V. et al. Imbalanced carbon-for-phosphorus exchange between European arbuscular mycorrhizal fungi and non-native Panicum grasses—a case of dysfunctional symbiosis. Pedobiologia 62, 48–55 (2017).
Article  Google Scholar 

72.
Ohno, T. & Zibilske, L. M. Determination of low concentrations of phosphorus in soil extracts using malachite green. Soil Sci. Soc. Am. J. 55, 892–895 (1991).
ADS  CAS  Article  Google Scholar 

73.
McGonigle, T. P., Miller, M. H., Evans, D. G., Fairchild, G. L. & Swan, J. A. A new method which gives an objective-measure of colonization of roots by vesicular arbuscular mycorrhizal fungi. New Phytol. 115, 495–501 (1990).
Article  Google Scholar 

74.
Koske, R. E. & Gemma, J. N. A modified procedure for staining roots to detect VA-mycorrhizas. Mycol. Res. 92, 486–505 (1989).
Article  Google Scholar 

75.
Gryndler, M. et al. Tuber aestivum Vittad. mycelium quantified: advantages and limitations of a qPCR approach. Mycorrhiza 23, 341–348 (2013).
PubMed  Article  PubMed Central  Google Scholar 

76.
Thonar, C., Erb, A. & Jansa, J. Real-time PCR to quantify composition of arbuscular mycorrhizal fungal communities-marker design, verification, calibration and field validation. Mol. Ecol. Res. 12, 219–232 (2012).
CAS  Article  Google Scholar 

77.
von Felten, A., Défago, G. & Maurhofer, M. Quantification of Pseudomonas fluorescens strains F113, CHA0 and Pf153 in the rhizosphere of maize by strain-specific real-time PCR unaffected by the variability of DNA extraction efficiency. J. Microbiol. Methods 81, 108–115 (2010).
Article  CAS  Google Scholar 

78.
Janoušková, M., Püschel, D., Hujslová, M., Slavíková, R. & Jansa, J. Quantification of arbuscular mycorrhizal fungal DNA in roots: how important is material preservation?. Mycorrhiza 25, 205–214 (2015).
PubMed  Article  CAS  Google Scholar  More

International Waterbird Census (IWC) data suggest 309,000 Scaup were wintering in North-West Europe in 1988–1991, compared with 192,300 in 2015–2018, indicating that the number of Scaup in this flyway has declined by 38.1% over 31 years (equivalent to a 30.3% decline over three generations, given a Scaup generation length of 8.2 years). Such a rate of decrease qualifies this population as vulnerable (VU) according to criterion A2(c) of the International Union for Conservation of Nature24. Thus, our results confirm the recent attribution of Scaup as a VU on the European Red List18. We suggest that the 1% threshold for the North-West Europe population of the Scaup should be revised to 1900.
In addition to the overall decline in abundance, we also show that changes in winter temperature on the eastern and northern edges of the wintering range potentially explain the observed dramatic shift in winter distribution closer to the breeding grounds. Climate change appears to have opened up more wintering sites to Scaup, especially in the more northern and eastern areas where reductions in winter ice cover have made previous staging sites increasingly accessible in winter. This might be expected to have a positive effect on the population, given that Scaup have more potential wintering sites to choose between and that they face a diminished risk from mass starvation because of the reduced probability of unexpected ice cover of potential feeding areas25. However, the ultimate causes of shifts in wintering distribution remain unknown and could equally relate to deterioration of food quality in southern and western wintering grounds. At Lake IJsselmeer, the annual changes in the large numbers of wintering Scaup there in the 1980s and 1990s were explained by fluctuations in the abundance of their main prey, Zebra Mussel Dreissena polymorpha26. The decline in Zebra Mussels in the IJsselmeer lake and its replacement by Quagga Mussels Dreissena rostriformis bugensis27 resulted in a deterioration in the quality of food resources at the site. These are likely contributory reasons to explain the shift in the centre of gravity of the Scaup wintering grounds to Poland and eastern Germany, although we lack data to determine the magnitude of this effect (Fig. 4, Unit#3). This area now constitutes the most important wintering area for this population, although the detection of Quagga Mussels in this region in 201428 represents a potential threat to the quality of this important wintering ground.
Assuming that some of the birds remain to winter along the migration route on sites formerly only used as stopovers, we can retrospectively infer the migration route of the Scaup population breeding in northern Russia and Fennoscandia (Fig. 1). It would appear that after birds reach the Baltic, they stop in Estonia before traversing the Baltic south-west to Gotland, migrating along the southern coast of Sweden and onwards to the main wintering area in Danish, German and Polish Baltic waters (Unit#3). Some Scaup continue west to reach Unit#2 in the Netherlands, and small numbers continue to reach France and the UK. The small population breeding in Iceland likely winter exclusively in the UK and Ireland, where fewer of the Russian/Fennoscandia population reach in recent winters. The Iceland breeding birds likely constitute a separate biogeographic population, with little contact with the main one discussed here (Fig. 1). Assuming the continuing effects of global warming, we can predict further separation of the two sub-populations and that Unit#4 (Fig. 4), the coast of Gotland and the islands and bays in Estonia will most likely play an increasingly important future role as winter quarters for this species. This is likely to be the case at other sites within eastern Baltic where this species can find suitable habitats.
Our historical analysis has shown that after a period of most rapid decline during 1988–2003, this population could be interpreted as remaining stable during 2003–2018 (Fig. 2). We suspect that this may be partly the result of the significant decrease in the Scaup bycatch in the Netherlands29,30,31. The added mortality from fisheries bycatch represents one of the most important threats to the relatively long-lived Scaup32. Evidence showed that drowning mortality was extremely high between 1985 and 1994, when an estimated average of 17,672 birds died annually in fishing gear (6% of the total population of the time), but this has declined since the 2000s32. Of all Scaup from this flyway population that drowned in fishing nets in years 1978–1990, up to 65% died at the most important wintering site at the time—the Dutch IJsselmeer32. However, our highly uneven knowledge of the extent of the Scaup bycatch throughout its winter range should be taken into account here. Exceptionally detailed estimates from IJsselmeer during the earlier period14 contrasts our lack of data or poor estimates from elsewhere, which may result in a bias that implies a greater importance for Scaup bycatch at the IJsselmeer for the population than was actually the case. Current estimates of bycatch levels throughout the flyway suggest that Scaup death in fishing nets has decreased, amounting to c.4000 individuals yearly, partly explained by the substantial decrease in the Dutch bycatch32.
The second highly important threat to Scaup, perhaps as important as the bycatch, is the deterioration of their food resources. Detailed energy budget studies on Lake IJsselmeer14 suggested that foraging Scaup there were operating on the margins of energetic profitability and the limited number of important wintering sites elsewhere suggest that alternative sites are really scarce, implying that food availability at core wintering sites could potentially affect winter survival.
The specialist habitat selection of the Scaup restricts it to a narrow range of habitats during the wintering period where it aggregates in large concentrations, a factor which causes the entire wintering population to concentrate in relatively few locations. Potentially, this makes them more vulnerable at the population level than most other, more dispersed diving duck species. During the January 2015 count, 91% of counted birds were present at 31 locations in five countries (Denmark, Germany, the Netherlands, Poland and Sweden). The four most important locations supported over two-thirds of the total wintering numbers: namely IJsselmeer in the Netherlands, Barther Bodden and Greifswalder Bodden in Germany and Odra river estuary in Poland (Fig. 4). Taken together, these areas have consistently been the most important wintering areas for Scaup over the last 30 years3,14,20, with two thirds of the flyway population during winter concentrated within 5300 km2 (2000 km2 in the Netherlands and 3300 km2 in Poland/Germany).
Wintering areas in Germany and Poland also act as stopover sites, so much larger numbers are counted there in autumn and spring migration, with up to 100,000 individuals on the Szczecin Lagoon (c.470 km29). Similarly, in Estonia, where a few hundred birds winter (Fig. 1), numbers may exceed 100,000 individuals in spring33. Therefore, cohesive planning for the effective conservation of the species, requires adequate protection at both the most important wintering sites (analysed in this article) and stopover sites along the entire migration route. During spring migration, extremely large Scaup concentrations can occur in these important sites, which provide for other biological functions such a communal courtship, displaying, pair-bonding etc.32. Given that Scaup are among the most vulnerable of diving ducks to bycatch34 (constituting more than 50% of diving birds drowned in fishing nets in the Polish Odra Estuary35) potentially high mortality during the prelude to the breeding season is likely to have severe adverse effects on the entire population. It is important to remember that this site can simultaneously support up to 75% of the total population9 and intensive fishing takes place here with gillnets35—the method of fishing recognised as the most dangerous for drowning diving birds in the Baltic Sea6.
Other environmental pressures on Scaup are no less serious, but currently less well quantified. Many important wintering areas are situated in estuaries of large rivers that invariably host major sea ports, where large vessels cause disturbance and pollution. Maintenance of shipping channels requires dredging (as in the case of the channel leading to the port of Amsterdam on IJsselmeer in the Netherlands and that serving the port of Szczecin on the Szczecin Lagoon in Poland). Dredging of shallow marine and brackish substrates can disrupt sediment horizons, mobilising suspended material, creating turbidity and disrupting the food resource and the ability of Scaup to forage for their prey. The proximity to human settlements also makes these shallow marine waters attractive to the increasing practice of water sports, kite- and wind-surfing, boating and recreational fishing from boats, which although not a source of direct mortality, contributes to disturbance and displacement of Scaup from favoured areas36,37.
SPAs and effectiveness of protection
The long-term conservation aim for a decreasing qualifying species, in accordance with European Union (EU) law (Birds Directive—Council Directive 2009/147/EC), should be to recover them to former level of abundance. To achieve this aim, SPAs should be designated in sites where 1% or more of the biogeographic population regularly occurs. In the case of Scaup, all of such areas are protected in the form of SPA (Table 2). Subsequently, such a SPAs should have a Management Plan (MP) defining the conservation objectives within each site, updated every 6 years. Of the three most important Scaup SPAs in Europe, only the IJsselmeer (NL9803028, Unit#2, Core wintering area, Fig. 4) has a MP for 2013–201738, which described the long term decline (since 1994) in wintering numbers of Scaup in the IJsselmeer and identified the greatest threats for Scaup as declining food resources and disturbance by developing water sports. Although bycatch was conspicuously not listed as a threat, the MP documents previous measures, taken to reduce fishing effort, had resulted from the implementation of another EU Directive—the Water Framework Directive (WFD, Directive 2000/60/EC). The WFD committed EU Member States to achieve good qualitative and quantitative status of all water bodies by 201538. Conservation measures carried out on Lake IJsselmeer over the last 75 years aimed to maintain sustainable fishing did not bring about the intended results on fish stock39. However, they may have had a positive effect on reducing bycatch of Scaup from 11,500 killed annually during 1978–199032 to insignificant numbers in the years 2011–201231, which may have contributed to the slowing in the rate of population decline at this time. In the most important wintering area for this flyway populations—the lagoons and bays either side of the German-Polish border, out of ten SPAs forming one coherent area (Fig. 4) only two have MPs. Moreover, the key SPAs within this area that regularly hold the highest Scaup numbers do not have MPs, they are: Greifswalder Bodden und südlicher Strelasund (DE1747402) in Germany and Szczecin Lagoon (PLB320009) in Poland. The Greifswalder Bodden, Teile des Strelasundes und Nordspitze Usedom (DE1747301) Special Area of Conservation (SAC), which overlaps with the Greifswalder Bodden und südlicher Strelasund SPA, was created under the Habitats Directive (Council Directive 92/43/EEC) and has a MP that identifies the threats to Scaup (e.g. from bycatch). However, because MPs for SACs (as against SPAs) are not primarily directed towards bird conservation, there are no specific regulations to limit the current stressors upon Scaup at this site40. The existing MPs for two other SPAs (“Vorpommersche Boddenlandschaft und nördlicher Strelasund” and “Dolina Dolnej Odry”) either do not identify main threats to Scaup or fail to impose sufficient conservation measures41,42.
Other SPAs that are less important for Scaup within Unit#3 west of the core wintering area include Östliche Kieler Bucht (DE1530491) and Ostsee östlich Wagrien (DE1633491), which have MPs identifying the threat from bycatch. This includes a voluntary agreement between the Schleswig Holstein Ministry of the Environment and local fishery associations, under which areas are closed to fishing if “concentrations of ducks” ( > 100) are present in the areas. Fishermen have two days to remove their gear after closure. There are rigid legal provisions at these two sites that prohibit fishing with gillnets within 200 m of the shore43,44. To date, there is no evidence of a positive effect and reduction of bycatch of diving birds, so we recommend a study of the effectiveness of these provisions.
The shift in the centre of gravity of the wintering population to Germany and Poland highlights the ineffectiveness of conservation measures directed towards Scaup (and other diving birds) there. Despite the existence of SPAs in which the Scaup is specifically protected and evidence of the cost of gillnet bycatch to local diving ducks, the most serious pressure remains unchecked. In 2011–2012, results from research work in the Szczecin Lagoon37 recommended the MP proposed reducing the Scaup (and other diving birds) bycatch by spatiotemporal regulation of gillnet fisheries to avoid key areas used by the birds. Unfortunately, the effective solutions to deliver results for bird conservation were considered too far-reaching by fishing interests. The fishing lobby blocked official approval of the MP by government and so these measures were never implemented. Given the high rates of Scaup bycatch, the designation of the area as a SPA offers no effective protection to the species at this site32. The effectiveness of SPA designation for a particular species remains ineffective, as long as effective management is not implemented. Given the increasing relative importance of the German/Polish resorts to the species in recent years, the lack of effective measures within these SPAs is becoming more critical to safeguard the conservation of the North-West Europe population of Scaup. Suitably prepared MPs, containing a bycatch monitoring order, would solve this problem, setting bycatch thresholds, according to the recommendations of BirdLife International45—1% of natural mortality calculated on the basis of local species abundance. If this value is exceeded, spatiotemporal restrictions on gillnet fishery should be introduced.
Looking to the future, areas that were formerly stopovers are already becoming wintering sites in Sweden and Estonia. Although currently not numerically significant in winter, these sites already hold significant numbers during migration. In the future, satellite areas (Unit#4) have the potential to develop into important wintering grounds and therefore require adequate protection from factors known to affect Scaup survival.
Previous studies show that bycatch in fishing nets is one of the most serious anthropogenic pressures during the non-breeding period for many diving birds6, although we cannot exclude the influence of other factors such as food availability and quality26 and disturbance from hunting46 and water sports37. Because the majority of North-West Europe’s Scaup winter in relatively few places, conservation interventions at these key sites are particularly important. The shift in wintering distribution poses new challenges for countries increasingly responsible for the conservation of this species in winter. Lack of adequate protection in this region means that these areas may act as sink habitats (in the sense of the source-sink model 47). The shift of wintering areas to sink habitats exposes an increasing part of the population to the pressures present there. This is not only the case for Scaup but also for a range of other diving bird species. These birds concentrate in the most attractive areas rich in food, often biologically productive transitional waters, where marine and freshwater birds meet in high densities. For this reason, effective conservation measures directed at Scaup will positively impact upon a whole range of other species with similar ecology. This suggests that protection measures taken for the Scaup could also benefit associated marine species in the same areas such as Long-tailed Duck, Velvet Scoter, Common Scoter Melanitta nigra, as well as for coastal zone species such as: Tufted Duck, Smew Mergellus albellus, and Goosander Mergus merganser. More

Adamo SA (2004) Estimating disease resistance in insects: phenoloxidase and lysozyme-like activity and disease resistance in the cricket Gryllus texensis. J Insect Physiol 50:209–216
CAS  PubMed  Article  Google Scholar 

Alcock J (1994) Postinsemination associations between males and females in insects: the mate-guarding hypothesis. Annu Rev Entomol 39:1–21
Article  Google Scholar 

Archer CR, Zajitschek F, Sakaluk SK, Royle NJ, Hunt J (2012) Sexual selection affects the evolution of lifespan and ageing in the decorated cricket Gryllodes sigillatus. Evolution 66:3088–3100
CAS  PubMed  Article  Google Scholar 

Arnqvist G (1988) Mate guarding and sperm displacement in the water strider Gerris lateralis Schumm. (Heteroptera: Gerridae). Freshw Biol 19:269–274
Article  Google Scholar 

Arnqvist G, Nilsson T (2000) The evolution of polyandry: multiple mating and female fitness in insects. Anim Behav 60:145–164
CAS  PubMed  Article  Google Scholar 

Arnqvist G, Rowe L (2002) Antagonistic coevolution between the sexes in a group of insects. Nature 415:787–789
CAS  PubMed  Article  Google Scholar 

Avila FW, Sirot LK, Laflamme BA, Rubinstein CD, Wolfner MF (2010) Insect seminal fluid proteins: Identification and function. Annu Rev Entomol 56:21–40
Article  CAS  Google Scholar 

Azad P, Ryu J, Haddad GG (2011) Distinct role of Hsp70 in Drosophila hemocytes during severe hypoxia. Free Radic Biol Med 51:530–538
CAS  PubMed  PubMed Central  Article  Google Scholar 

Baer B, Morgan ED, Schmid-Hempel P (2001) A nonspecific fatty acid within the bumblebee mating plug prevents females from remating. Proc Natl Acad Sci U S A 98:3926–3928
CAS  PubMed  PubMed Central  Article  Google Scholar 

Barribeau SM, Schmid-Hempel P (2017) Sexual healing: mating induces a protective immune response in bumblebees. J Evol Biol 30:202–209
CAS  PubMed  Article  Google Scholar 

Bascuñán-García AP, Lara C, Córdoba-Aguilar A (2010) Immune investment impairs growth, female reproduction and survival in the house cricket, Acheta domesticus. J Insect Physiol 56:204–211
PubMed  Article  CAS  Google Scholar 

Bates D, Mächler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1–48.
Article  Google Scholar 

Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach, 2nd edn. Springer-Verlag, New York
Google Scholar 

Burpee DM, Sakaluk SK (1993) Repeated matings offset costs of reproduction in female crickets. Evol Ecol 7:240–250
Article  Google Scholar 

Castella G, Christe P, Chapuisat (2009) Mating triggers dynamic immune regulations in wood ant queen. J Evol Biol 22:564–570
CAS  PubMed  Article  Google Scholar 

Chapman T, Arnqvist G, Bangham J, Rowe L (2003) Sexual conflict. Trends Ecol Evol 18:41–47
Article  Google Scholar 

Cordero A (1990) The adaptive significance of the prolonged copulations of the damselfly, Ischnura graellsii (Odonata: Coenagrionidae). Anim Behav 40:43–48
Article  Google Scholar 

Cordero A (1999) Forced copulations and female contact guarding at a high male density in a calopterygid damselfly. J Insect Behav 12:27–37
Article  Google Scholar 

Delbare SYN, Chow CY, Wolfner MF, Clark AG (2017) Roles of female and male genotype in post-mating responses in Drosophila melanogaster. J Hered 4:740–753
Article  CAS  Google Scholar 

Dickinson JL, Rutowski RL (1989) The function of the mating plug in the chalcedon checkerspot butterfly. Anim Behav 38:154–162
Article  Google Scholar 

Dougherty LR, van Lieshout E, McNamara KB, Moschilla JA, Arnqvist G, Simmons LW (2017) Sexual conflict and correlated evolution between male persistence and female resistance traits in the seed beetle Callosobruchus maculatus. Proc R Soc B Biol Sci 284:20170132
Article  Google Scholar 

Duffield KR, Hampton KJ, Houslay TM, Hunt J, Rapkin J, Sakaluk SK, Sadd BM (2018) Age-dependent variation in the terminal investment threshold in male crickets. Evolution 72:578–589
PubMed  Article  Google Scholar 

Duffield KR, Hampton KJ, Houslay TM, Hunt J, Sadd BM, Sakaluk SK (2019) Inbreeding alters context‐dependent reproductive effort and immunity in male crickets. J Evol Biol 32:731–741
PubMed  Google Scholar 

Edward DA, Poissant J, Wilson AJ, Chapman T (2014) Sexual conflict and interacting phenotypes: a quantitative genetic analysis of fecundity and copula duration in Drosophila melanogaster. Evolution 68:1651–1660
PubMed  Article  Google Scholar 

Eggert AK, Reinhardt K, Sakaluk SK (2003) Linear models for assessing mechanisms of sperm competition: the trouble with transformations. Evolution 57:173–176
PubMed  Article  Google Scholar 

Fedorka KM, Zuk M (2005) Sexual conflict and female immune suppression in the cricket, Allonemobious socius. J Evol Biol 18:1515–1522
PubMed  Article  Google Scholar 

Fedorka KM, Zuk M, Mousseau TA (2004) Immune suppression and the cost of reproduction in the ground cricket, Allonemobius socius. Evolution 58:2478–2485
PubMed  Article  Google Scholar 

Fedorka KM, Linder JE, Winterhalter W, Promislow D (2007) Post-mating disparity between potential and realized immune response in Drosophila melanogaster. Proc R Soc B Biol Sci 274:1211–1217
Article  Google Scholar 

Fricke C, Ávila‐Calero S, Armitage SA (2020) Genotypes and their interaction effects on reproduction and mating‐induced immune activation in Drosophila melanogaster. J Evol Biol 33:930–941
CAS  PubMed  Article  Google Scholar 

Gershman SN, Barnett CA, Pettinger AM, Weddle CB, Hunt J, Sakaluk SK (2010a) Inbred decorated crickets exhibit higher measures of macroparasitic immunity than outbred individuals. Heredity 105:282–289
CAS  PubMed  Article  Google Scholar 

Gershman SN, Barnett CA, Pettinger AM, Weddle CB, Hunt J, Sakaluk SK (2010b) Give ‘til it hurts: trade-offs between immunity and male reproductive effort in the decorated cricket, Gryllodes sigillatus. J Evol Biol 23:829–839
CAS  PubMed  Article  Google Scholar 

Gershman SN, Hunt J, Sakaluk SK (2013) Food fight: sexual conflict over free amino acids in the nuptial gifts of male decorated crickets. J Evol Biol 26:693–704
CAS  PubMed  Article  Google Scholar 

Gershman SN, Mitchell C, Sakaluk SK, Hunt J (2012) Biting off more than you can chew: sexual selection on the free amino acid composition of the spermatophylax in decorated crickets. Proc R Soc B Biol Sci 279:2531–2538
CAS  Article  Google Scholar 

Gillespie JP, Kanost MR, Trenczek T (1997) Biological mediators of insect immunity. Annu Rev Entomol 42:611–654
CAS  PubMed  Article  Google Scholar 

Gillott C (2003) Male accessory gland secretions: modulators of female reproductive physiology and behavior. Annu Rev Entomol 48:163–184
CAS  PubMed  Article  Google Scholar 

Goenaga J, Yamane T, Rönn J, Arnqvist G (2015) Within-species divergence in the seminal fluid proteome and its effect on male and female reproduction in a beetle. BMC Evol Biol 15:266
PubMed  PubMed Central  Article  CAS  Google Scholar 

González-Santoyo I, Córdoba-Aguilar A (2012) Phenoloxidase: a key component of the insect immune system. Entomol Exp Appl 142:1–16
Article  CAS  Google Scholar 

Haerty W, Jagadeeshan S, Kulathinal RJ, Wong A, Ravi Ram K, Sirot LK et al (2007) Evolution in the fast lane: Rapidly evolving sex-related genes in Drosophila. Genetics 177:1321–1335
CAS  PubMed  PubMed Central  Article  Google Scholar 

Hall MD, Lailvaux SP, Brooks RC (2013) Sex‐specific evolutionary potential of pre‐and postcopulatory reproductive interactions in the field cricket, Teleogryllus commodus. Evolution 67:1831–1837
PubMed  Article  PubMed Central  Google Scholar 

Hardin JW, Hilbe, JM (2007) Generalized linear models and extensions. 2nd edn, Stata Press, College Station, Texas

Ivy TM, Sakaluk SK (2005) Polyandry promotes enhanced offspring survival in decorated crickets. Evolution 59:152–159
PubMed  Article  Google Scholar 

Ivy TM, Weddle CB, Sakaluk SK (2005) Females use self-referent cues to avoid mating with previous mates. Proc R Soc B Biol Sci 272:2475–2478
Article  Google Scholar 

Kacsoh BZ, Schlenke TA (2012) High hemocyte load is associated with increased resistance against parasitoids in Drosophila suzukii, a relative of D. melanogaster. PLoS ONE 7:e34721
CAS  PubMed  PubMed Central  Article  Google Scholar 

Kerr AM, Gershman SN, Sakaluk SK (2010) Experimentally induced spermatophore production and immune responses reveal a trade-off in crickets. Behav Ecol 21:647–654
Article  Google Scholar 

Klowden MJ (1999) The check is in the male: Male mosquitoes affect female physiology and behavior. J Am Mosq Contr 15:213–220
CAS  Google Scholar 

Kwon H, Smith RC (2019) Chemical depletion of phagocytic immune cells in Anopheles gambiae reveals dual roles of mosquito hemocytes in anti-Plasmodium immunity. Proc Natl Acad Sci U S A 116:14119–14128
CAS  PubMed  PubMed Central  Article  Google Scholar 

Lawniczak MKN, Barnes AI, Linklater JR, Boone JM, Wigby S, Chapman T (2007) Mating and immunity in invertebrates. Trends Ecol Evol 22:48–55
PubMed  Article  Google Scholar 

Lavine MD, Strand MR (2002) Insect hemocytes and their role in immunity. Insect Biochem Mol Biol 32:1295–1309
CAS  PubMed  Article  Google Scholar 

Lenth, SingmannH, Love J, Buerkner P, Herve M (2020) emmeans: estimated marginal means, aka least-squares means. Release 1.4.5. https://CRAN.R-project.org/package=emmeans

Lung O, Wolfner MF (2001) Identification and characterization of the major Drosophila melanogaster mating plug protein. Insect Biochem Mol Biol 31:543–551
CAS  PubMed  Article  Google Scholar 

Miller JS, Nguyen T, Stanley-Samuelson DW (1994) Eicosanoids mediate insect nodulation responses to bacterial infections. Proc Natl Acad Sci U S A 91:12418–12422
CAS  PubMed  PubMed Central  Article  Google Scholar 

Morrow EH, Innocenti P (2012) Female postmating immune responses, immune system evolution and immunogenic males. Biol Rev 87:631–638
PubMed  Article  Google Scholar 

Nappi AJ, Vass E (1993) Melanogenesis and the generation of cytotoxic molecules during insect cellular immune reactions. Pigment Cell Res 6:117–126
CAS  PubMed  Article  Google Scholar 

Oku K, Price TAR, Wedell N (2019) Does mating negatively affect female immune defences in insects? Anim Biol 69:117–136
Article  Google Scholar 

Otti O (2015) Genitalia‐associated microbes in insects. Insect Sci 22:325–339
PubMed  Article  Google Scholar 

Parker GA, Birkhead TR (2013) Polyandry: the history of a revolution. Philos Trans R Soc Lond B Biol Sci 368:1–13
Article  Google Scholar 

Pauchet Y, Wielsch N, Wilkinson PA, Sakaluk SK, Svatoš A, ffrench-Constant RH et al (2015) What’s in the gift? Towards a molecular dissection of nuptial feeding in a cricket. PLoS ONE 10:e0140191
PubMed  PubMed Central  Article  CAS  Google Scholar 

Peng J, Zipperlen P, Kubli E (2005) Drosophila sex-peptide stimulates female innate immune system after mating via the Toll and Imd pathways. Curr Biol 15:1690–1694
CAS  PubMed  Article  Google Scholar 

Perry JC, Siro L, Wigby S (2013) The seminal symphony: how to compose an ejaculate. Trends Ecol Evol 28:414–422
PubMed  PubMed Central  Article  Google Scholar 

Ramirez JL, Garver LS, Brayner FA, Alves LC, Rodrigues J, Molina-Cruz A et al (2014) The role of hemocytes in Anopheles gambiae antiplasmodial immunity. J Innate Immun 6:119–128
CAS  PubMed  Article  Google Scholar 

Ravi Ram K, Wolfner MF (2007) Seminal influences: Drosophila acps and the molecular interplay between males and females during reproduction. Integr Comp Biol 47:427–445
CAS  PubMed  Article  Google Scholar 

R Core Team (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria

Rice WR, Holland B (1997) The enemies within: intergenomic conflict, interlocus contest evolution (ICE), and the intraspecific red queen. Behav Ecol Sociobiol 41:1–10
Article  Google Scholar 

Rolff J, Siva-Jothy MT (2002) Copulation corrupts immunity: a mechanism for a cost of mating in insects. Proc Natl Acad Sci U S A 99:9916–9918
CAS  PubMed  PubMed Central  Article  Google Scholar 

Rowe L, Arnqvist G (2002) Sexually antagonistic coevolution in a mating system: cominbing experimental and comparative approaches to address evolutionary processes. Evolution 56:754–767
PubMed  Article  Google Scholar 

Sakaluk SK (1984) Male crickets feed females to ensure complete sperm transfer. Science 223:609–610
CAS  PubMed  Article  Google Scholar 

Sakaluk SK (1986) Sperm competition and the evolution of nuptial feeding behavior in the cricket, Gryllodes supplicans (Walker). Evolution 40:584–593
PubMed  Article  Google Scholar 

Sakaluk SK (1987) Reproductive behaviour of the decorated cricket, Gryllodes supplicans (Orthoptera: Gryllidae): calling schedules, spatial distribution, and mating. Behaviour 100:202–225
Article  Google Scholar 

Sakaluk SK (1991) Post-copulatory mate guarding in decorated crickets. Anim Behav 41:207–216
Article  Google Scholar 

Sakaluk SK (2000) Sensory exploitation as an evolutionary origin to nuptial food gifts in insects. Proc R Soc Lond B Biol Sci 267:339–343

Sakaluk SK, Avery RL, Weddle CB (2006) Cryptic sexual conflict in gift-giving insects: chasing the chase-away. Am Nat 167:94–104
PubMed  Article  Google Scholar 

Sakaluk SK, Duffield KR, Rapkin J, Sadd BM, Hunt J (2019) The troublesome gift: the spermatophylax as a purveyor of sexual conflict and coercion in crickets. Adv Stud Behav 51:1–30
Article  Google Scholar 

Sakaluk SK, Eggert AK (1996) Female control of sperm transfer and intraspecific variation in sperm precedence: antecedents to the evolution of a courtship food gift. Evolution 50:694–703
PubMed  Article  Google Scholar 

Sakaluk SK, Schaus JM, Eggert AK, Snedden WA, Brady PL (2002) Polyandry and fitness of offspring reared under varying nutritional stress in decorated crickets. Evolution 56:1999–2007
PubMed  Article  Google Scholar 

Schneider CA, Rasband WS, Eliceiri KW (2012) NIH Image to ImageJ: 25 years of image analysis. Nat Methods 9:671–675
CAS  PubMed  PubMed Central  Article  Google Scholar 

Schneider PM (1985) Purification and properties of three lysozymes from hemolymph of the cricket, Gryllus bimaculatus (De Geer). Insect Biochem 15:463–470
CAS  Article  Google Scholar 

Schwenke RA, Lazzaro BP, Wolfner MF (2016) Reproduction-immunity trade-offs in insects. Annu Rev Entomol 61:239–256
CAS  PubMed  Article  Google Scholar 

Sherman KJ (1983) The adaptive significance of postcopulatory mate guarding in a dragonfly, Pachydiplax longipennis. Anim Behav 31:1107–1115
Article  Google Scholar 

Shoemaker KL, Parsons NM, Adamo SA (2006) Mating enhances parasite resistance in the cricket Gryllus texensis. Anim Behav 71:371–380
Article  Google Scholar 

Short SM, Lazzaro BP (2010) Female and male genetic contributions to post-mating immune defence in female Drosophila melanogaster. Proc R Soc B Biol Sci 277:3649–3657
Article  Google Scholar 

Soderhall K, Cerenius L (1998) Role of the prophenoloxidase-activating system in invertebrate immunity. Curr Opin Immunol 10:23–28
CAS  PubMed  Article  Google Scholar 

Stanley D, Kim Y (2014) Eicosanoid signaling in insects: from discovery to plant protection. Crit Rev Plant Sci 33:20–63
CAS  Article  Google Scholar 

Sugumaran M, Nellaiappan K, Valivittan K (2000) A new mechanism for the control of phenoloxidase activity: inhibition and complex formation with quinone isomerase. Arch Biochem Biophys 379:252–260
CAS  PubMed  Article  Google Scholar 

Theopold U, Schmidt O, Söderhäll K, Dushay MS (2004) Coagulation in arthropods: defence, wound closure and healing. Trends Immunol 25:289–294
CAS  PubMed  Article  Google Scholar 

Therneau T (2020) A package for survival analysis in R. Release 3.1-11. https://CRAN.R-project.org/package=survival

Warwick S, Vahed K, Raubenheimer D, Simpson SJ (2009) Free amino acids as phagostimulants in cricket nuptial gifts: support for the “candymaker” hypothesis. Biol Lett 5:194–196
CAS  PubMed  PubMed Central  Article  Google Scholar 

Wigby S, Chapman T (2005) Sex peptide causes mating costs in female Drosophila melanogaster. Curr Biol 15:316–321
CAS  PubMed  Article  Google Scholar 

Wolfner MF (1997) Tokens of love: functions and regulation of Drosophila male accessory gland products. Insect Biochem Mol Biol 27:179–192
CAS  PubMed  Article  Google Scholar 

Worthington AM, Jurenka RA, Kelly CD (2015) Mating for male-derived prostaglandin: a functional explanation for the increased fecundity of mated female crickets? J Exp Biol 218:2720–2727
PubMed  Article  Google Scholar 

Worthington AM, Kelly CD (2016) Females gain survival benefits from immune-boosting ejaculates. Evolution 70:928–933
PubMed  Article  Google Scholar 

Yi HY, Chowdhury M, Huang YD, Yu XQ (2014) Insect antimicrobial peptides and their applications. Appl Microbiol Biotechnol 98:5807–5822
CAS  PubMed  PubMed Central  Article  Google Scholar 

Zhong W, Priest NK, McClure CD, Evans CR, Mlynski DT, Immonen E et al (2013) Immune anticipation of mating in Drosophila: turandot m promotes immunity against sexually transmitted fungal infections. Proc R Soc B Biol Sci 280:1–9
Google Scholar  More