Rands, M. R. W. et al. Biodiversity conservation: challenges beyond 2010. Science 329, 1298–1303 (2010).
Brook, B. W., Sodhi, N. S. & Bradshaw, C. J. A. Synergies among extinction drivers under global change. Trends Ecol. Evol. 23, 453–460 (2008).
Butchart, S. H. M. et al. Global biodiversity: indicators of recent declines. Science 328, 1164–1168 (2010).
Crooks, K. R. et al. Quantification of habitat fragmentation reveals extinction risk in terrestrial mammals. Proc. Natl Acad. Sci. USA 114, 7635–7640 (2017).
Haddad, N. M. et al. Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci. Adv. 1, e1500052 (2015).
Charlesworth, D. & Willis, J. H. The genetics of inbreeding depression. Nat. Rev. Genet. 10, 783–796 (2009).
Kardos, M. et al. The crucial role of genome-wide genetic variation in conservation. Proc. Natl Acad. Sci. USA 118, e2104642118 (2021).
Lynch, M., Conery, J. & Burger, R. Mutation accumulation and the extinction of small populations. Am. Nat. 146, 489–518 (1995).
Dussex, N., Morales, H. E., Grossen, C., Dalén, L. & van Oosterhout, C. Purging and accumulation of genetic load in conservation. Trends Ecol. Evol. 38, 961–969 (2023).
Keller, L. F. & Waller, D. M. Inbreeding effects in wild populations. Trends Ecol. Evol. 17, 230–241 (2002).
Bijlsma, R. & Loeschcke, V. Genetic erosion impedes adaptive responses to stressful environments. Evol. Appl. 5, 117–129 (2012).
Reed, D. H. & Frankham, R. Correlation between fitness and genetic diversity. Conserv. Biol. 17, 230–237 (2003).
Willi, Y., Van Buskirk, J. & Hoffmann, A. A. Limits to the adaptive potential of small populations. Annu. Rev. Ecol. Evol. Syst. 37, 433–458 (2006).
Bell, D. A. et al. The exciting potential and remaining uncertainties of genetic rescue. Trends Ecol. Evol. 34, 1070–1079 (2019).
Whiteley, A. R., Fitzpatrick, S. W., Funk, W. C. & Tallmon, D. A. Genetic rescue to the rescue. Trends Ecol. Evol. 30, 42–49 (2015).
Tallmon, D. A., Luikart, G. & Waples, R. S. The alluring simplicity and complex reality of genetic rescue. Trends Ecol. Evol. 19, 489–496 (2004).
Gilpin, M. E. & Soulé, M. E. in Conservation Biology: the Science of Scarcity and Diversity (ed. Soulé, M. E.) 19–34 (Sinauer Associates, 1986).
Weeks, A. R. et al. Genetic rescue increases fitness and aids rapid recovery of an endangered marsupial population. Nat. Commun. 8, 1071 (2017).
Vilà, C. et al. Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant. Proc. R. Soc. Lond. B 270, 91–97 (2003).
Johnson, W. E. et al. Genetic restoration of the Florida panther. Science 329, 1641–1645 (2010).
Fitzpatrick, S. W. et al. Genomic and fitness consequences of genetic rescue in wild populations. Curr. Biol. 30, 517–522.e5 (2020).
Miller, S. M. et al. Genetic rescue of an isolated African lion population. Conserv. Genet. 21, 41–53 (2020).
Madsen, T. et al. Genetic rescue restores long-term viability of an isolated population of adders (Vipera berus). Curr. Biol. 30, R1297–R1299 (2020).
Pregler, K. C. et al. Assisted gene flow from outcrossing shows the potential for genetic rescue in an endangered salmon population. Conserv. Lett. 16, e12934 (2023).
Hogg, J. T., Forbes, S. H., Steele, B. M. & Luikart, G. Genetic rescue of an insular population of large mammals. Proc. R. Soc. Lond. B 273, 1491–1499 (2006).
Nichols, S. et al. Genetic rescue attempt in a small, inbred population of a wild endangered passerine. Biol. Conserv. 290, 110430 (2024).
Pavlova, A. et al. Planning and implementing genetic rescue of an endangered freshwater fish population in a regulated river, where low flow reduces breeding opportunities and may trigger inbreeding depression. Evol. Appl. 17, e13679 (2024).
Clarke, J. G., Smith, A. C. & Cullingham, C. I. Genetic rescue often leads to higher fitness as a result of increased heterozygosity across animal taxa. Mol. Ecol. 33, e17532 (2024).
Frankham, R. Genetic rescue benefits persist to at least the F3 generation, based on a meta-analysis. Biol. Conserv. 195, 33–36 (2016).
Edmands, S. Between a rock and a hard place: evaluating the relative risks of inbreeding and outbreeding for conservation and management. Mol. Ecol. 16, 463–475 (2007).
MacDonald, Z. G. et al. Whole-genome evaluation of genetic rescue: the case of a curiously isolated and endangered butterfly. Mol. Ecol. 34, e17657 (2025).
Kyriazis, C. C., Wayne, R. K. & Lohmueller, K. E. Strongly deleterious mutations are a primary determinant of extinction risk due to inbreeding depression. Evol. Lett. 5, 33–47 (2021).
Frankham, R., Ballou, J. D. & Briscoe, D. A. Introduction to Conservation Genetics (Cambridge Univ. Press, 2010).
Whitlock, R. et al. A systematic review of phenotypic responses to between-population outbreeding. Environ. Evid. 2, 13 (2013).
Robinson, J. A., Brown, C., Kim, B. Y., Lohmueller, K. E. & Wayne, R. K. Purging of strongly deleterious mutations explains long-term persistence and absence of inbreeding depression in island foxes. Curr. Biol. 28, 3487–3494 (2018).
Hedrick, P. W., Robinson, J. A., Peterson, R. O. & Vucetich, J. A. Genetics and extinction and the example of Isle Royale wolves. Anim. Conserv. 22, 302–309 (2019).
Hedrick, P. W., Peterson, R. O., Vucetich, L. M., Adams, J. R. & Vucetich, J. A. Genetic rescue in Isle Royale wolves: genetic analysis and the collapse of the population. Conserv. Genet. 15, 1111–1121 (2014).
Korona, R. Genetic load of the yeast Saccharomyces cerevisiae under diverse environmental conditions. Evolution 53, 1966–1971 (1999).
Kondrashov, A. S. & Houle, D. Genotype—environment interactions and the estimation of the genomic mutation rate in Drosophila melanogaster. Proc. R. Soc. Lond. B 258, 221–227 (1994).
Heber, S., Briskie, J. V. & Apiolaza, L. A. A test of the ‘genetic rescue’ technique using bottlenecked donor populations of Drosophila melanogaster. PLoS ONE 7, e43113 (2012).
Pérez-Pereira, N., Caballero, A. & García-Dorado, A. Reviewing the consequences of genetic purging on the success of rescue programs. Conserv. Genet. 23, 1–17 (2022).
Zajitschek, S. R., Zajitschek, F. & Brooks, R. C. Demographic costs of inbreeding revealed by sex-specific genetic rescue effects. BMC Evol. Biol. 9, 289 (2009).
West, G. et al. Sexual selection matters in genetic rescue, but productivity benefits fade over time: a multi-generation experiment to inform conservation. Proc. R. Soc. Lond. B 292, 20242374 (2025).
Bateman, A. J. Intra-sexual selection in Drosophila. Heredity 2, 349–368 (1948).
Andersson, M. Sexual Selection (Princeton Univ. Press, 1994).
Rowe, L. & Houle, D. The lek paradox and the capture of genetic variance by condition dependent traits. Proc. R. Soc. Lond. B 263, 1415–1421 (1996).
Lumley, A. J. et al. Sexual selection protects against extinction. Nature 522, 470–473 (2015).
Bonduriansky, R. & Chenoweth, S. F. Intralocus sexual conflict. Trends Ecol. Evol. 24, 280–288 (2009).
Plesnar-Bielak, A., Skrzynecka, A. M., Miler, K. & Radwan, J. Selection for alternative male reproductive tactics alters intralocus sexual conflict. Evolution 68, 2137–2144 (2014).
Harano, T., Okada, K., Nakayama, S., Miyatake, T. & Hosken, D. J. Intralocus sexual conflict unresolved by sex-limited trait expression. Curr. Biol. 20, 2036–2039 (2010).
Whitlock, M. C. & Agrawal, A. F. Purging the genome with sexual selection: reducing mutation load through selection on males. Evolution 63, 569–582 (2009).
Dugand, R. J., Tomkins, J. L. & Kennington, W. J. Molecular evidence supports a genic capture resolution of the lek paradox. Nat. Commun. 10, 1359 (2019).
Parrett, J. M. et al. Genomic evidence that a sexually selected trait captures genome-wide variation and facilitates the purging of genetic load. Nat. Ecol. Evol. 6, 1330–1342 (2022).
Parrett, J. M. & Knell, R. J. The effect of sexual selection on adaptation and extinction under increasing temperatures. Proc. R. Soc. Lond. B 285, 20180303 (2018).
Jarzebowska, M. & Radwan, J. Sexual selection counteracts extinction of small populations of the bulb mites. Evolution 64, 1283–1289 (2010).
Ralls, K. et al. Call for a paradigm shift in the genetic management of fragmented populations. Conserv. Lett. 11, e12412 (2018).
Pérez-Pereira, N., Kleinman-Ruiz, D., García-Dorado, A., Quesada, H. & Caballero, A. A test of the long-term efficiency of genetic rescue with Drosophila melanogaster. Mol. Ecol. 34, e17690 (2025).
Ralls, K., Sunnucks, P., Lacy, R. C. & Frankham, R. Genetic rescue: a critique of the evidence supports maximizing genetic diversity rather than minimizing the introduction of putatively harmful genetic variation. Biol. Conserv. 251, 108784 (2020).
Lynch, M. et al. Perspective: spontaneous deleterious mutation. Evolution 53, 645–663 (1999).
Haldane, J. B. S. The effect of variation of fitness. Am. Nat. 71, 337–349 (1937).
Chen, R. S., Soulsbury, C. D., Hench, K., van Oers, K. & Hoffman, J. I. Predicted deleterious mutations reveal the genetic architecture of male reproductive success in a lekking bird. Nat. Ecol. Evol. 9, 1924–1937 (2025).
Mezmouk, S. & Ross-Ibarra, J. The pattern and distribution of deleterious mutations in maize. G3: Genes Genomes Genet. 4, 163–171 (2014).
Kryukov, G. V., Pennacchio, L. A. & Sunyaev, S. R. Most rare missense alleles are deleterious in humans: implications for complex disease and association studies. Am. J. Hum. Genet. 80, 727–739 (2007).
Łukasiewicz, A., Niśkiewicz, M. & Radwan, J. Sexually selected male weapon is associated with lower inbreeding load but higher sex load in the bulb mite. Evolution 74, 1851–1855 (2020).
Joag, R. et al. Transcriptomics of intralocus sexual conflict: gene expression patterns in females change in response to selection on a male secondary sexual trait in the bulb mite. Genome Biol. Evol. 8, 2351–2357 (2016).
Parrett, J. M., Sobala, K., Chmielewski, S., Przesmycka, K. & Radwan, J. No evidence of negative frequency-dependent selection in alternative reproductive tactics in a bulb mite. Anim. Behav. 220, 123048 (2025).
Radwan, J. & Klimas, M. Male dimorphism in the bulb mite, Rhizoglyphus robini: fighters survive better. Ethol. Ecol. Evol. 13, 69–79 (2001).
Radwan, J., Czyz, M., Konior, M. & Kołodziejczyk, M. Aggressiveness in two male morphs of the bulb mite Rhizoglyphus robini. Ethology 106, 53–62 (2000).
Plesnar-Bielak, A., Jawor, A. & Kramarz, P. E. Complex response in size-related traits of bulb mites (Rhizoglyphus robini) under elevated thermal conditions – an experimental evolution approach. J. Exp. Biol. 216, 4542–4548 (2013).
Parrett, J. M., Kulczak, M. & Szudarek-Trepto, N. Fertility loss under thermal stress: males have lower fertility limits but no evidence of sex differences in sensitivity. Oikos 2024, e10329 (2024).
Radwan, J. Male morph determination in two species of acarid mites. Heredity 74, 669–673 (1995).
Parrett, J. M. et al. A sexually selected male weapon characterized by strong additive genetic variance and no evidence for sexually antagonistic polyphenic maintenance. Evolution 77, 1289–1302 (2023).
Smallegange, I. M. Complex environmental effects on the expression of alternative reproductive phenotypes in the bulb mite. Evol. Ecol. 25, 857–873 (2011).
Rhebergen, F. T., Stewart, K. A. & Smallegange, I. M. Nutrient-dependent allometric plasticity in a male-diphenic mite. Ecol. Evol. 12, e9145 (2022).
Łukasiewicz, A., Porwal, N., Niśkiewicz, M., Parrett, J. M. & Radwan, J. Sexually selected male weapon increases the risk of population extinction under environmental change: an experimental evidence. Evolution 77, 2291–2300 (2023).
Radwan, J. Inbreeding depression in fecundity and inbred line extinction in the bulb mite, Rhizoglyphus robini. Heredity 90, 371–376 (2003).
Spielman, D., Brook, B. W. & Frankham, R. Most species are not driven to extinction before genetic factors impact them. Proc. Natl Acad. Sci. USA 101, 15261–15264 (2004).
Radwan, J. & Siva-Jothy, M. T. The function of post-insemination mate association in the bulb mite, Rhizoglyphus robini. Anim. Behav. 52, 651–657 (1996).
Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014).
Chmielewski, S. et al. Sex-specific recombination landscape in a species with holocentric chromosomes. Genetics 231, iyaf217 (2025).
Li, H. Aligning sequence reads, clone sequences and assembly contigs with BWA-MEM. Preprint at https://arxiv.org/abs/1303.3997 (2013).
Li, H. et al. The Sequence Alignment/Map format and SAMtools. Bioinformatics 25, 2078–2079 (2009).
Kofler, R., Pandey, R. V. & Schlötterer, C. PoPoolation2: identifying differentiation between populations using sequencing of pooled DNA samples (Pool-Seq). Bioinformatics 27, 3435–3436 (2011).
Kofler, R. et al. Popoolation: a toolbox for population genetic analysis of next generation sequencing data from pooled individuals. PLoS ONE 6, e15925 (2011).
Plesnar-Bielak, A. et al. Transcriptomics of differences in thermal plasticity associated with selection for an exaggerated male sexual trait. Heredity 133, 43–53 (2024).
Li, H. A statistical framework for SNP calling, mutation discovery, association mapping and population genetical parameter estimation from sequencing data. Bioinformatics 27, 2987–2993 (2011).
Quinlan, A. R. & Hall, I. M. BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics 26, 841–842 (2010).
Cingolani, P. et al. A program for annotating and predicting the effects of single nucleotide polymorphisms, SnpEff: SNPs in the genome of Drosophila melanogaster strain w1118; iso-2; iso-3. Fly (Austin) 6, 80–92 (2012).
R Core Team R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2024).
Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer-Verlag, 2016).
Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R Journal 9, 378–400 (2017).
Therneau, T. M. coxme: mixed effects Cox models. R Package version 2.2-22 https://cran.r-project.org/web/packages/coxme/index.html (2024).
Parrett, J. Genetic rescue increases long-term fitness despite elevating putative genetic load in a male-dimorphic mite. figshare https://doi.org/10.6084/m9.figshare.30165784 (2026).