{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T22:46:04Z","timestamp":1791931564163,"version":"4.3.4"},"reference-count":118,"publisher":"Springer Science and Business Media LLC","issue":"5","license":[{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T00:00:00Z","timestamp":1778112000000},"content-version":"vor","delay-in-days":6,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100006447","name":"Universit\u00e4t Z\u00fcrich","doi-asserted-by":"publisher","award":["FK-20-088"],"award-info":[{"award-number":["FK-20-088"]}],"id":[{"id":"10.13039\/501100006447","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001711","name":"Schweizerischer Nationalfonds zur F\u00f6rderung der Wissenschaftlichen Forschung","doi-asserted-by":"publisher","award":["310030_197651"],"award-info":[{"award-number":["310030_197651"]}],"id":[{"id":"10.13039\/501100001711","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Behav Ecol Sociobiol"],"published-print":{"date-parts":[[2026,5]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Because reproductive tactics vary in their energetic and fitness costs, individuals may increase their reproductive success by aligning their tactics with their physical condition. In resource-based mating systems, monopolizing valuable patches is energetically costly, yet potentially highly rewarding for males. In females, mating near resources may reduce the cost of mate search and assessment. However, the balance of costs and benefits associated with a given tactic may shift with an individual\u2019s condition. Here we studied this in the fruit fly\n                    <jats:italic>Drosophila prolongata<\/jats:italic>\n                    . In arenas containing resource-dense areas (RDAs) typically used as territories, we tracked the individual positions and behaviors of flies in groups using machine-learning-based pose estimation and behavioral classification. We found that large, high-condition males achieved elevated mating success through territoriality and courtship, whilst small males obtained copulations through high mobility, consistent with a scramble competition tactic. Female behavior was also condition dependent: large females spent more time in RDAs, mated sooner at those sites, and thus reinforced territory-based sexual selection, whereas small females mated later and more frequently outside RDAs, potentially avoiding harassment costs. In addition, we detected size-disassortative pairings, suggesting that the complex cost-benefit interactions encountered by females of varying condition can lead to counter-intuitive associations. However, given the limited duration of our observations and the high remating rate in this species, the prevalence of this disassortative mating pattern over longer time scales warrants further investigation. Our findings reveal how resource distribution and individual condition may interact to maintain multiple reproductive tactics in both sexes.\n                  <\/jats:p>","DOI":"10.1007\/s00265-026-03739-z","type":"journal-article","created":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T11:06:43Z","timestamp":1778152003000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Condition-dependent success of resource-based reproductive tactics across both sexes in Drosophila prolongata"],"prefix":"10.1007","volume":"80","author":[{"given":"Alessio N.","family":"De Nardo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Swarnima","family":"Mukherjee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefan","family":"L\u00fcpold","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,7]]},"reference":[{"key":"3739_CR1","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1007\/s002650050638","volume":"46","author":"DJ Able","year":"1999","unstructured":"Able DJ (1999) Scramble competition selects for greater tailfin size in male red-spotted newts (Amphibia: Salamandridae). Behav Ecol Sociobiol 46:423\u2013428. https:\/\/doi.org\/10.1007\/s002650050638","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR2","doi-asserted-by":"publisher","first-page":"210","DOI":"10.2108\/zs200010","volume":"37","author":"K Amino","year":"2020","unstructured":"Amino K, Matsuo T (2020) Intra- versus inter-sexual selection on sexually dimorphic traits in Drosophila prolongata. Zoolog Sci 37:210. https:\/\/doi.org\/10.2108\/zs200010","journal-title":"Zoolog Sci"},{"key":"3739_CR3","doi-asserted-by":"publisher","first-page":"104831","DOI":"10.1016\/j.beproc.2023.104831","volume":"206","author":"K Amino","year":"2023","unstructured":"Amino K, Matsuo T (2023) Reproductive advantage of the winners of male-male competition in Drosophila prolongata. Behav Processes 206:104831. https:\/\/doi.org\/10.1016\/j.beproc.2023.104831","journal-title":"Behav Processes"},{"key":"3739_CR4","doi-asserted-by":"publisher","DOI":"10.1515\/9780691207278","volume-title":"Sexual selection","author":"M Andersson","year":"1994","unstructured":"Andersson M (1994) Sexual selection. Princeton University Press, Princeton (N.J.)"},{"key":"3739_CR5","doi-asserted-by":"publisher","first-page":"296","DOI":"10.1016\/j.tree.2006.03.015","volume":"21","author":"M Andersson","year":"2006","unstructured":"Andersson M, Simmons LW (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296\u2013302. https:\/\/doi.org\/10.1016\/j.tree.2006.03.015","journal-title":"Trends Ecol Evol"},{"key":"3739_CR6","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1006\/anbe.2000.1617","volume":"61","author":"PW Bateman","year":"2001","unstructured":"Bateman PW, Gilson LN, Ferguson JWH (2001) Male size and sequential mate preference in the cricket Gryllus bimaculatus. Anim Behav 61:631\u2013637. https:\/\/doi.org\/10.1006\/anbe.2000.1617","journal-title":"Anim Behav"},{"key":"3739_CR7","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v067.i01","volume":"67","author":"D Bates","year":"2015","unstructured":"Bates D, M\u00e4chler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1\u201348. https:\/\/doi.org\/10.18637\/jss.v067.i01","journal-title":"J Stat Softw"},{"key":"3739_CR8","doi-asserted-by":"publisher","first-page":"7898","DOI":"10.1038\/s41598-020-64941-1","volume":"10","author":"M Belenioti","year":"2020","unstructured":"Belenioti M, Chaniotakis N (2020) Aggressive behaviour of Drosophila suzukii in relation to environmental and social factors. Sci Rep 10:7898. https:\/\/doi.org\/10.1038\/s41598-020-64941-1","journal-title":"Sci Rep"},{"key":"3739_CR9","first-page":"247","volume":"57","author":"M Berec","year":"2011","unstructured":"Berec M, Bajgar A (2011) Choosy outsiders? Satellite males associate with sexy hosts in the European tree frog Hyla arborea. Acta Zool Acad Sci Hung 57:247\u2013254","journal-title":"Acta Zool Acad Sci Hung"},{"key":"3739_CR10","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1007\/BF01237760","volume":"9","author":"WU Blanckenhorn","year":"1995","unstructured":"Blanckenhorn WU, Preziosi RF, Fairbairn DJ (1995) Time and energy constraints and the evolution of sexual size dimorphism \u2014 to eat or to mate? Evol Ecol 9:369\u2013381. https:\/\/doi.org\/10.1007\/BF01237760","journal-title":"Evol Ecol"},{"key":"3739_CR11","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1111\/j.1439-0310.2011.01889.x","volume":"117","author":"LA Bonachea","year":"2011","unstructured":"Bonachea LA, Ryan MJ (2011) Simulated predation risk influences female choice in T\u00fangara frogs, Physalaemus pustulosus. Ethology 117:400\u2013407. https:\/\/doi.org\/10.1111\/j.1439-0310.2011.01889.x","journal-title":"Ethology"},{"key":"3739_CR12","doi-asserted-by":"publisher","first-page":"1385","DOI":"10.1007\/s10682-011-9523-5","volume":"25","author":"J Bro-J\u00f8rgensen","year":"2011","unstructured":"Bro-J\u00f8rgensen J (2011) Queuing in space and time reduces the lek paradox on an antelope lek. Evol Ecol 25:1385\u20131395. https:\/\/doi.org\/10.1007\/s10682-011-9523-5","journal-title":"Evol Ecol"},{"key":"3739_CR13","first-page":"160","volume":"76","author":"JL Brown","year":"1964","unstructured":"Brown JL (1964) The evolution of diversity in avian territorial systems. Wilson Bull 76:160\u2013169","journal-title":"Wilson Bull"},{"key":"3739_CR14","doi-asserted-by":"crossref","unstructured":"Buzzatto BA, Tomkins JL, Simmons LW (2014) Alternative phenotypes within mating systems. In: Shuker D, Simmons LW (eds) The Evolution of Insect Mating Systems. Oxford University Press, Oxford","DOI":"10.1093\/acprof:oso\/9780199678020.003.0007"},{"key":"3739_CR15","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1086\/497401","volume":"166","author":"JA Byers","year":"2005","unstructured":"Byers JA, Wiseman PA, Jones L, Roffe TJ (2005) A large cost of female mate sampling in pronghorn. Am Nat 166:661\u2013668. https:\/\/doi.org\/10.1086\/497401","journal-title":"Am Nat"},{"key":"3739_CR16","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1046\/j.1365-2656.2002.00619.x","volume":"71","author":"R Calsbeek","year":"2002","unstructured":"Calsbeek R, Sinervo B (2002) An experimental test of the ideal despotic distribution. J Anim Ecol 71:513\u2013523. https:\/\/doi.org\/10.1046\/j.1365-2656.2002.00619.x","journal-title":"J Anim Ecol"},{"key":"3739_CR17","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1034\/j.1600-0706.2001.950204.x","volume":"95","author":"U Candolin","year":"2001","unstructured":"Candolin U, Voigt H-R (2001) Correlation between male size and territory quality: consequence of male competition or predation susceptibility? Oikos 95:225\u2013230. https:\/\/doi.org\/10.1034\/j.1600-0706.2001.950204.x","journal-title":"Oikos"},{"key":"3739_CR18","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1016\/S0003-3472(05)80667-0","volume":"40","author":"J Carranza","year":"1990","unstructured":"Carranza J, Alvarez F, Redondo T (1990) Territoriality as a mating strategy in red deer. Anim Behav 40:79\u201388. https:\/\/doi.org\/10.1016\/S0003-3472(05)80667-0","journal-title":"Anim Behav"},{"key":"3739_CR19","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1111\/mec.12577","volume":"23","author":"BA Caspers","year":"2014","unstructured":"Caspers BA, Krause ET, Hendrix R et al (2014) The more the better \u2013 polyandry and genetic similarity are positively linked to reproductive success in a natural population of terrestrial salamanders (Salamandra salamandra). Mol Ecol 23:239\u2013250. https:\/\/doi.org\/10.1111\/mec.12577","journal-title":"Mol Ecol"},{"key":"3739_CR20","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/j.anbehav.2008.09.017","volume":"77","author":"S Castellano","year":"2009","unstructured":"Castellano S (2009) Unreliable preferences, reliable choice and sexual selection in leks. Anim Behav 77:225\u2013232. https:\/\/doi.org\/10.1016\/j.anbehav.2008.09.017","journal-title":"Anim Behav"},{"key":"3739_CR21","doi-asserted-by":"publisher","first-page":"5664","DOI":"10.1073\/pnas.082102599","volume":"99","author":"S Chen","year":"2002","unstructured":"Chen S, Lee AY, Bowens NM et al (2002) Fighting fruit flies: A model system for the study of aggression. Proc Natl Acad Sci 99:5664\u20135668. https:\/\/doi.org\/10.1073\/pnas.082102599","journal-title":"Proc Natl Acad Sci"},{"key":"3739_CR23","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1007\/BF00166699","volume":"30","author":"S Dale","year":"1992","unstructured":"Dale S, Rinden H, Slagsvold T (1992) Competition for a mate restricts mate search of female pied flycatchers. Behav Ecol Sociobiol 30:165\u2013176. https:\/\/doi.org\/10.1007\/BF00166699","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR22","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1111\/brv.13129","volume":"100","author":"AV Da Silva","year":"2025","unstructured":"Da Silva AV, De Almeida JGL, Ventura SPR et al (2025) A meta-analysis on alternative mating tactics: when the main and the alternative yield similar reproductive success. Biol Rev 100:85\u201398. https:\/\/doi.org\/10.1111\/brv.13129","journal-title":"Biol Rev"},{"key":"3739_CR27","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1093\/beheco\/aru072","volume":"25","author":"R Deb","year":"2014","unstructured":"Deb R, Balakrishnan R (2014) The opportunity for sampling: the ecological context of female mate choice. Behav Ecol 25:967\u2013974. https:\/\/doi.org\/10.1093\/beheco\/aru072","journal-title":"Behav Ecol"},{"key":"3739_CR28","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.anbehav.2016.02.019","volume":"114","author":"F-X Dechaume-Moncharmont","year":"2016","unstructured":"Dechaume-Moncharmont F-X, Brom T, C\u00e9zilly F (2016) Opportunity costs resulting from scramble competition within the choosy sex severely impair mate choosiness. Anim Behav 114:249\u2013260. https:\/\/doi.org\/10.1016\/j.anbehav.2016.02.019","journal-title":"Anim Behav"},{"key":"3739_CR24","doi-asserted-by":"publisher","first-page":"20242501","DOI":"10.1098\/rspb.2024.2501","volume":"292","author":"AN De Nardo","year":"2025","unstructured":"De Nardo AN, Biswas B, Perdig\u00f3n Ferreira J et al (2025a) Socio-ecological context modulates the significance of territorial contest competition in Drosophila prolongata. Proc R Soc B Biol Sci 292:20242501. https:\/\/doi.org\/10.1098\/rspb.2024.2501","journal-title":"Proc R Soc B Biol Sci"},{"key":"3739_CR25","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1093\/evolut\/qpaf035","volume":"79","author":"AN De Nardo","year":"2025","unstructured":"De Nardo AN, Meena A, Maggu K et al (2025b) Experimental evolution reveals trade-offs between sexual selection and heat tolerance in Drosophila prolongata. Evolution 79:823\u2013836. https:\/\/doi.org\/10.1093\/evolut\/qpaf035","journal-title":"Evolution"},{"key":"3739_CR26","doi-asserted-by":"publisher","first-page":"2014","DOI":"10.1111\/evo.14228","volume":"75","author":"AN De Nardo","year":"2021","unstructured":"De Nardo AN, Roy J, Sbilordo SH, L\u00fcpold S (2021) Condition-dependent interaction between mating success and competitive fertilization success in Drosophila melanogaster. Evolution 75:2014\u20132026. https:\/\/doi.org\/10.1111\/evo.14228","journal-title":"Evolution"},{"key":"3739_CR29","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1007\/s00265-002-0555-2","volume":"53","author":"CE deRivera","year":"2003","unstructured":"deRivera CE, Backwell PRY, Christy JH, Vehrencamp SL (2003) Density affects female and male mate searching in the fiddler crab, Uca beebei. Behav Ecol Sociobiol 53:72\u201383","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR30","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1007\/s00265-007-0500-5","volume":"62","author":"PD Dijkstra","year":"2008","unstructured":"Dijkstra PD, van der Zee EM, Groothuis TGG (2008) Territory quality affects female preference in a Lake Victoria cichlid fish. Behav Ecol Sociobiol 62:747\u2013755. https:\/\/doi.org\/10.1007\/s00265-007-0500-5","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR31","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1093\/beheco\/arac100","volume":"34","author":"LR Dougherty","year":"2023","unstructured":"Dougherty LR (2023) The effect of individual state on the strength of mate choice in females and males. Behav Ecol 34:197\u2013209. https:\/\/doi.org\/10.1093\/beheco\/arac100","journal-title":"Behav Ecol"},{"key":"3739_CR32","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1093\/beheco\/14.1.2","volume":"14","author":"F Dubois","year":"2003","unstructured":"Dubois F, Giraldeau L-A, Grant JWA (2003) Resource defense in a group-foraging context. Behav Ecol 14:2\u20139. https:\/\/doi.org\/10.1093\/beheco\/14.1.2","journal-title":"Behav Ecol"},{"key":"3739_CR33","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1038\/s41598-020-57481-1","volume":"10","author":"C Edomwande","year":"2020","unstructured":"Edomwande C, Barbosa F (2020) The influence of predation risk on mate signaling and mate choice in the lesser waxmoth Achroia grisella. Sci Rep 10:524. https:\/\/doi.org\/10.1038\/s41598-020-57481-1","journal-title":"Sci Rep"},{"key":"3739_CR34","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1126\/science.327542","volume":"197","author":"ST Emlen","year":"1977","unstructured":"Emlen ST, Oring LW (1977) Ecology, sexual selection, and the evolution of mating systems. Science 197:215\u2013223. https:\/\/doi.org\/10.1126\/science.327542","journal-title":"Science"},{"key":"3739_CR35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.tpb.2014.06.001","volume":"96","author":"AM Frame","year":"2014","unstructured":"Frame AM, Mills AF (2014) Condition-dependent mate choice: A stochastic dynamic programming approach. Theor Popul Biol 96:1\u20137. https:\/\/doi.org\/10.1016\/j.tpb.2014.06.001","journal-title":"Theor Popul Biol"},{"key":"3739_CR36","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1093\/beheco\/8.2.162","volume":"8","author":"CR Gabor","year":"1997","unstructured":"Gabor CR, Halliday TR (1997) Sequential mate choice by multiply mating smooth newts: females become more choosy. Behav Ecol 8:162\u2013166. https:\/\/doi.org\/10.1093\/beheco\/8.2.162","journal-title":"Behav Ecol"},{"key":"3739_CR37","volume-title":"The Economy of nature and the evolution of sex","author":"MT Ghiselin","year":"1974","unstructured":"Ghiselin MT (1974) The Economy of nature and the evolution of sex. University of California Press, Berkeley, Los Angeles, London"},{"key":"3739_CR38","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1016\/S0003-3472(05)80922-4","volume":"40","author":"LM Gosling","year":"1990","unstructured":"Gosling LM, Petrie M (1990) Lekking in topi: a consequence of satellite behaviour by small males at hotspots. Anim Behav 40:272\u2013287. https:\/\/doi.org\/10.1016\/S0003-3472(05)80922-4","journal-title":"Anim Behav"},{"key":"3739_CR39","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1080\/10236249309378862","volume":"23","author":"JWA Grant","year":"1993","unstructured":"Grant JWA (1993) Whether or not to defend? The influence of resource distribution. Mar Behav Physiol 23:137\u2013153. https:\/\/doi.org\/10.1080\/10236249309378862","journal-title":"Mar Behav Physiol"},{"key":"3739_CR40","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/0169-5347(96)81050-0","volume":"11","author":"MR Gross","year":"1996","unstructured":"Gross MR (1996) Alternative reproductive strategies and tactics: diversity within sexes. Trends Ecol Evol 11:92\u201398. https:\/\/doi.org\/10.1016\/0169-5347(96)81050-0","journal-title":"Trends Ecol Evol"},{"key":"3739_CR41","doi-asserted-by":"publisher","first-page":"626","DOI":"10.1111\/j.1439-0310.2005.01086.x","volume":"111","author":"PA Guerra","year":"2005","unstructured":"Guerra PA, Mason AC (2005) Information on resource quality mediates aggression between male Madagascar hissing cockroaches, Gromphadorhina portentosa (Dictyoptera: Blaberidae). Ethology 111:626\u2013637. https:\/\/doi.org\/10.1111\/j.1439-0310.2005.01086.x","journal-title":"Ethology"},{"key":"3739_CR42","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1038\/346172a0","volume":"346","author":"DT Gwynne","year":"1990","unstructured":"Gwynne DT, Simmons LW (1990) Experimental reversal of courtship roles in an insect. Nature 346:172\u2013174. https:\/\/doi.org\/10.1038\/346172a0","journal-title":"Nature"},{"key":"3739_CR43","doi-asserted-by":"publisher","first-page":"488","DOI":"10.1111\/j.1439-0310.1997.tb00162.x","volume":"103","author":"MA Hack","year":"1997","unstructured":"Hack MA, Thompson DJ, Fernandes DM (1997) Fighting in males of the autumn spider, Metellina segmentata: effects of relative body size, prior residency and female value on contest outcome and duration. Ethology 103:488\u2013498. https:\/\/doi.org\/10.1111\/j.1439-0310.1997.tb00162.x","journal-title":"Ethology"},{"key":"3739_CR44","doi-asserted-by":"publisher","first-page":"320","DOI":"10.1111\/jeb.12563","volume":"28","author":"T Harano","year":"2015","unstructured":"Harano T (2015) Receptive females mitigate costs of sexual conflict. J Evol Biol 28:320\u2013327. https:\/\/doi.org\/10.1111\/jeb.12563","journal-title":"J Evol Biol"},{"key":"3739_CR45","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1007\/s10164-012-0331-3","volume":"30","author":"T Harano","year":"2012","unstructured":"Harano T, Sato N, Miyatake T (2012) Effects of female and male size on female mating and remating decisions in a bean beetle. J Ethol 30:337\u2013343. https:\/\/doi.org\/10.1007\/s10164-012-0331-3","journal-title":"J Ethol"},{"key":"3739_CR46","doi-asserted-by":"crossref","unstructured":"Hartig F (2016) DHARMa: residual diagnostics for hierarchical (multi-level \/ mixed) regression models. 0.4.7","DOI":"10.32614\/CRAN.package.DHARMa"},{"key":"3739_CR47","doi-asserted-by":"crossref","unstructured":"Herberstein ME, Painting CJ, Holwell GI (2017) Scramble competition polygyny in terrestrial arthropods. Advances in the Study of Behavior. Elsevier, Cambridge, pp 237\u2013295","DOI":"10.1016\/bs.asb.2017.01.001"},{"key":"3739_CR48","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1016\/S0003-3472(05)80533-0","volume":"40","author":"AA Hoffmann","year":"1990","unstructured":"Hoffmann AA, Cacoyianni Z (1990) Territoriality in Drosophila melanogaster as a conditional strategy. Anim Behav 40:526\u2013537. https:\/\/doi.org\/10.1016\/S0003-3472(05)80533-0","journal-title":"Anim Behav"},{"key":"3739_CR49","doi-asserted-by":"publisher","DOI":"10.1086\/430672","author":"J Hunt","year":"2005","unstructured":"Hunt J, Brooks R, Jennions MD (2005) Female mate choice as a condition-dependent life\u2010history trait. Am Nat. https:\/\/doi.org\/10.1086\/430672","journal-title":"Am Nat"},{"key":"3739_CR50","unstructured":"Iannone R (2023) DiagrammeR: graph\/network visualization"},{"key":"3739_CR51","doi-asserted-by":"publisher","first-page":"e1500983","DOI":"10.1126\/sciadv.1500983","volume":"2","author":"T Janicke","year":"2016","unstructured":"Janicke T, H\u00e4derer IK, Lajeunesse MJ, Anthes N (2016) Darwinian sex roles confirmed across the animal kingdom. Sci Adv 2:e1500983. https:\/\/doi.org\/10.1126\/sciadv.1500983","journal-title":"Sci Adv"},{"key":"3739_CR52","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1017\/S0006323196005014","volume":"72","author":"MD Jennions","year":"1997","unstructured":"Jennions MD, Petrie M (1997) Variation in mate choice and mating preferences: a review of causes and consequences. Biol Rev Camb Philos Soc 72:283\u2013327. https:\/\/doi.org\/10.1017\/S0006323196005014","journal-title":"Biol Rev Camb Philos Soc"},{"key":"3739_CR53","doi-asserted-by":"publisher","first-page":"29","DOI":"10.4161\/fly.5.1.13713","volume":"5","author":"T Jonsson","year":"2011","unstructured":"Jonsson T, Kravitz EA, Heinrich R (2011) Sound production during agonistic behavior of male Drosophila melanogaster. Fly (Austin) 5:29\u201338. https:\/\/doi.org\/10.4161\/fly.5.1.13713","journal-title":"Fly (Austin)"},{"key":"3739_CR54","doi-asserted-by":"publisher","first-page":"906","DOI":"10.1093\/beheco\/art017","volume":"24","author":"AT Kahn","year":"2013","unstructured":"Kahn AT, Dolstra T, Jennions MD, Backwell PRY (2013) Strategic male courtship effort varies in concert with adaptive shifts in female mating preferences. Behav Ecol 24:906\u2013913. https:\/\/doi.org\/10.1093\/beheco\/art017","journal-title":"Behav Ecol"},{"key":"3739_CR55","doi-asserted-by":"publisher","first-page":"1073","DOI":"10.1093\/beheco\/araa054","volume":"31","author":"CD Kelly","year":"2020","unstructured":"Kelly CD (2020) Sexual selection on size and shape in Japanese beetles (Popillia japonica). Behav Ecol 31:1073\u20131083. https:\/\/doi.org\/10.1093\/beheco\/araa054","journal-title":"Behav Ecol"},{"key":"3739_CR56","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1086\/589894","volume":"172","author":"CD Kelly","year":"2008","unstructured":"Kelly CD, Bussi\u00e8re LF, Gwynne DT (2008) Sexual selection for male mobility in a giant insect with female-biased size dimorphism. Am Nat 172:417\u2013423. https:\/\/doi.org\/10.1086\/589894","journal-title":"Am Nat"},{"key":"3739_CR57","doi-asserted-by":"publisher","first-page":"e422","DOI":"10.1371\/journal.pone.0000422","volume":"2","author":"TW Kim","year":"2007","unstructured":"Kim TW, Christy JH, Choe JC (2007) A preference for a sexual signal keeps females safe. PLoS ONE 2:e422. https:\/\/doi.org\/10.1371\/journal.pone.0000422","journal-title":"PLoS ONE"},{"key":"3739_CR58","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1111\/j.1469-7998.2009.00566.x","volume":"278","author":"RJ Knell","year":"2009","unstructured":"Knell RJ (2009) Population density and the evolution of male aggression. J Zool 278:83\u201390. https:\/\/doi.org\/10.1111\/j.1469-7998.2009.00566.x","journal-title":"J Zool"},{"key":"3739_CR59","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1007\/BF00300545","volume":"19","author":"A Kodric-Brown","year":"1986","unstructured":"Kodric-Brown A (1986) Satellites and sneakers: opportunistic male breeding tactics in pupfish (Cyprinodon pecosensis). Behav Ecol Sociobiol 19:425\u2013432. https:\/\/doi.org\/10.1007\/BF00300545","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR60","doi-asserted-by":"publisher","first-page":"1876","DOI":"10.1111\/j.0014-3820.2005.tb01058.x","volume":"59","author":"H Kokko","year":"2005","unstructured":"Kokko H, Mappes J (2005) Sexual selection when fertilization is not guaranteed. Evolution 59:1876\u20131885. https:\/\/doi.org\/10.1111\/j.0014-3820.2005.tb01058.x","journal-title":"Evolution"},{"key":"3739_CR61","doi-asserted-by":"publisher","first-page":"2109","DOI":"10.1098\/rspb.1999.0895","volume":"266","author":"H Kokko","year":"1999","unstructured":"Kokko H, Rintamaki PT, Alatalo RV et al (1999) Female choice selects for lifetime lekking performance in black grouse males. Proc R Soc Lond B Biol Sci 266:2109\u20132115. https:\/\/doi.org\/10.1098\/rspb.1999.0895","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"3739_CR62","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1007\/s00265-022-03172-y","volume":"76","author":"V Kovalov","year":"2022","unstructured":"Kovalov V, Kokko H (2022) Selection for male stamina can help explain costly displays with cost-minimizing female choice. Behav Ecol Sociobiol 76:69. https:\/\/doi.org\/10.1007\/s00265-022-03172-y","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR63","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1111\/ens.12116","volume":"18","author":"A Kudo","year":"2015","unstructured":"Kudo A, Takamori H, Watabe H et al (2015) Variation in morphological and behavioral traits among isofemale strains of Drosophila prolongata (Diptera: Drosophilidae). Entomol Sci 18:221\u2013229. https:\/\/doi.org\/10.1111\/ens.12116","journal-title":"Entomol Sci"},{"key":"3739_CR64","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/beheco\/8.3.268","volume":"8","author":"RB Lanctot","year":"1997","unstructured":"Lanctot RB, Weatherhead PJ (1997) Ephemeral lekking behavior in the buff-breasted sandpiper, Tryngites subruficollis. Behav Ecol 8:268\u2013278. https:\/\/doi.org\/10.1093\/beheco\/8.3.268","journal-title":"Behav Ecol"},{"key":"3739_CR69","doi-asserted-by":"publisher","first-page":"772","DOI":"10.21105\/joss.00772","volume":"3","author":"D L\u00fcdecke","year":"2018","unstructured":"L\u00fcdecke D (2018) ggeffects: tidy data frames of marginal effects from regression models. J Open Source Softw 3:772. https:\/\/doi.org\/10.21105\/joss.00772","journal-title":"J Open Source Softw"},{"key":"3739_CR65","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1111\/2041-210X.12512","volume":"7","author":"JS Lefcheck","year":"2016","unstructured":"Lefcheck JS (2016) piecewiseSEM: Piecewise structural equation modelling in r for ecology, evolution, and systematics. Methods Ecol Evol 7:573\u2013579. https:\/\/doi.org\/10.1111\/2041-210X.12512","journal-title":"Methods Ecol Evol"},{"key":"3739_CR66","unstructured":"Lenth RV (2023) emmeans: Estimated Marginal Means. aka Least-Square Means"},{"key":"3739_CR67","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1139\/z90-092","volume":"68","author":"SL Lima","year":"1990","unstructured":"Lima SL, Dill LM (1990) Behavioral decisions made under the risk of predation: a review and prospectus. Can J Zool 68:619\u2013640. https:\/\/doi.org\/10.1139\/z90-092","journal-title":"Can J Zool"},{"key":"3739_CR68","doi-asserted-by":"publisher","first-page":"20130983","DOI":"10.1098\/rspb.2013.0983","volume":"280","author":"K Lindstr\u00f6m","year":"2013","unstructured":"Lindstr\u00f6m K, Lehtonen TK (2013) Mate sampling and choosiness in the sand goby. Proc R Soc B Biol Sci 280:20130983. https:\/\/doi.org\/10.1098\/rspb.2013.0983","journal-title":"Proc R Soc B Biol Sci"},{"key":"3739_CR70","doi-asserted-by":"publisher","first-page":"1686","DOI":"10.1111\/evo.13246","volume":"71","author":"S L\u00fcpold","year":"2017","unstructured":"L\u00fcpold S, Jin L, Liao WB (2017) Population density and structure drive differential investment in pre- and postmating sexual traits in frogs. Evolution 71:1686\u20131699. https:\/\/doi.org\/10.1111\/evo.13246","journal-title":"Evolution"},{"key":"3739_CR71","doi-asserted-by":"publisher","first-page":"20180443","DOI":"10.1098\/rsbl.2018.0443","volume":"14","author":"A MacPherson","year":"2018","unstructured":"MacPherson A, Yun L, Barrera TS et al (2018) The effects of male harm vary with female quality and environmental complexity in Drosophila melanogaster. Biol Lett 14:20180443. https:\/\/doi.org\/10.1098\/rsbl.2018.0443","journal-title":"Biol Lett"},{"key":"3739_CR72","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1111\/evo.13455","volume":"72","author":"EL McCullough","year":"2018","unstructured":"McCullough EL, Buzatto BA, Simmons LW (2018) Population density mediates the interaction between pre- and postmating sexual selection. Evolution 72:893\u2013905. https:\/\/doi.org\/10.1111\/evo.13455","journal-title":"Evolution"},{"key":"3739_CR73","doi-asserted-by":"publisher","unstructured":"McLain DK (1992) Population density and the intensity of sexual selection on body length in spatially or temporally restricted natural populations of a seed bug. Behav Ecol Sociobiol 30. https:\/\/doi.org\/10.1007\/BF00170602","DOI":"10.1007\/BF00170602"},{"key":"3739_CR74","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1007\/s00265-003-0616-1","volume":"54","author":"SC Mills","year":"2003","unstructured":"Mills SC, Reynolds JD (2003) Operational sex ratio and alternative reproductive behaviours in the European bitterling, Rhodeus sericeus. Behav Ecol Sociobiol 54:98\u2013104. https:\/\/doi.org\/10.1007\/s00265-003-0616-1","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR75","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.anbehav.2018.04.002","volume":"140","author":"K Minekawa","year":"2018","unstructured":"Minekawa K, Miyatake T, Ishikawa Y, Matsuo T (2018) The adaptive role of a species-specific courtship behaviour in coping with remating suppression of mated females. Anim Behav 140:29\u201337. https:\/\/doi.org\/10.1016\/j.anbehav.2018.04.002","journal-title":"Anim Behav"},{"key":"3739_CR76","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1098\/rsbl.2007.0300","volume":"3","author":"J Moya-Lara\u00f1o","year":"2007","unstructured":"Moya-Lara\u00f1o J, El-Sayyid MET, Fox CW (2007) Smaller beetles are better scramble competitors at cooler temperatures. Biol Lett 3:475\u2013478. https:\/\/doi.org\/10.1098\/rsbl.2007.0300","journal-title":"Biol Lett"},{"key":"3739_CR77","doi-asserted-by":"publisher","unstructured":"Mukherjee S, De Nardo AN, L\u00fcpold S (2026) Sex-specific site preferences in response to resource value in Drosophila prolongata. Behav Ecol Sociobiol 80:63. https:\/\/doi.org\/10.1007\/s00265-026-03731-7","DOI":"10.1007\/s00265-026-03731-7"},{"key":"3739_CR78","unstructured":"Muro A (2021) Examining female mate choice based on relatedness or size of males in Drosophila prolongata. MSc thesis, University of Zurich"},{"key":"3739_CR79","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1007\/s00442-021-05068-6","volume":"197","author":"TJ Ord","year":"2021","unstructured":"Ord TJ (2021) Costs of territoriality: a review of hypotheses, meta-analysis, and field study. Oecologia 197:615\u2013631. https:\/\/doi.org\/10.1007\/s00442-021-05068-6","journal-title":"Oecologia"},{"key":"3739_CR80","unstructured":"Parker GA (1983) Mate quality and mating decisions. In: Bateson P (ed) Mate choice. Cambridge University Press, pp 141\u2013164"},{"key":"3739_CR81","doi-asserted-by":"publisher","first-page":"1269","DOI":"10.1111\/j.1558-5646.1994.tb05311.x","volume":"48","author":"L Partridge","year":"1994","unstructured":"Partridge L, Barrie B, Fowler K, French V (1994) Evolution and development of body size and cell size in Drosophila melanogaster in response to temperature. Evolution 48:1269\u20131276. https:\/\/doi.org\/10.1111\/j.1558-5646.1994.tb05311.x","journal-title":"Evolution"},{"key":"3739_CR82","first-page":"2825","volume":"12","author":"F Pedregosa","year":"2011","unstructured":"Pedregosa F, Varoquaux G, Gramfort A et al (2011) Scikit-learn: machine learning in Python. J Mach Learn Res 12:2825\u20132830","journal-title":"J Mach Learn Res"},{"key":"3739_CR83","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1093\/beheco\/arab127","volume":"33","author":"J Perdig\u00f3n Ferreira","year":"2022","unstructured":"Perdig\u00f3n Ferreira J, L\u00fcpold S (2022) Condition- and context-dependent alternative reproductive tactic in Drosophila prolongata. Behav Ecol 33:213\u2013221. https:\/\/doi.org\/10.1093\/beheco\/arab127","journal-title":"Behav Ecol"},{"key":"3739_CR84","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1038\/s41592-022-01426-1","volume":"19","author":"TD Pereira","year":"2022","unstructured":"Pereira TD, Tabris N, Matsliah A et al (2022) SLEAP: A deep learning system for multi-animal pose tracking. Nat Methods 19:486\u2013495. https:\/\/doi.org\/10.1038\/s41592-022-01426-1","journal-title":"Nat Methods"},{"key":"3739_CR85","doi-asserted-by":"publisher","first-page":"1623","DOI":"10.1098\/rspb.2002.2280","volume":"270","author":"TE Pitcher","year":"2003","unstructured":"Pitcher TE, Neff BD, Rodd FH, Rowe L (2003) Multiple mating and sequential mate choice in guppies: females trade up. Proc R Soc Lond B Biol Sci 270:1623\u20131629. https:\/\/doi.org\/10.1098\/rspb.2002.2280","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"3739_CR86","doi-asserted-by":"publisher","first-page":"1821","DOI":"10.1098\/rspb.2002.2090","volume":"269","author":"S Pitnick","year":"2002","unstructured":"Pitnick S, Garc\u00eda\u2013Gonz\u00e1lez F (2002) Harm to females increases with male body size in Drosophila melanogaster. Proc R Soc Lond B Biol Sci 269:1821\u20131828. https:\/\/doi.org\/10.1098\/rspb.2002.2090","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"3739_CR87","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/S0022-5193(87)80169-8","volume":"128","author":"A Pomiankowski","year":"1987","unstructured":"Pomiankowski A (1987) The costs of choice in sexual selection. J Theor Biol 128:195\u2013218","journal-title":"J Theor Biol"},{"key":"3739_CR89","doi-asserted-by":"publisher","first-page":"1851","DOI":"10.1016\/j.anbehav.2007.11.002","volume":"75","author":"G Raihani","year":"2008","unstructured":"Raihani G, Serrano-Meneses MA, C\u00f3rdoba-Aguilar A (2008) Male mating tactics in the American rubyspot damselfly: territoriality, nonterritoriality and switching behaviour. Anim Behav 75:1851\u20131860. https:\/\/doi.org\/10.1016\/j.anbehav.2007.11.002","journal-title":"Anim Behav"},{"key":"3739_CR90","doi-asserted-by":"publisher","first-page":"569","DOI":"10.1007\/s10905-015-9528-y","volume":"28","author":"J Ram\u00edrez-Delgado","year":"2015","unstructured":"Ram\u00edrez-Delgado J, L\u00f3pez-Garc\u00eda K, Lara C, Serrano-Meneses MA (2015) Wing pigmentation in males of a territorial damselfly: alternative reproductive tactics, allometry and mating success. J Insect Behav 28:569\u2013581. https:\/\/doi.org\/10.1007\/s10905-015-9528-y","journal-title":"J Insect Behav"},{"key":"3739_CR88","unstructured":"R Core Team (2023) R: A language and environment for statistical computing"},{"key":"3739_CR91","doi-asserted-by":"publisher","first-page":"1569","DOI":"10.1111\/j.1365-294X.2004.02151.x","volume":"13","author":"M Reichard","year":"2004","unstructured":"Reichard M, Smith C, Jordan WC (2004) Genetic evidence reveals density-dependent mediated success of alternative mating behaviours in the European bitterling (Rhodeus sericeus). Mol Ecol 13:1569\u20131578. https:\/\/doi.org\/10.1111\/j.1365-294X.2004.02151.x","journal-title":"Mol Ecol"},{"key":"3739_CR92","doi-asserted-by":"publisher","first-page":"1707","DOI":"10.1111\/ecog.03382","volume":"41","author":"PT Rohner","year":"2018","unstructured":"Rohner PT, Pitnick S, Blanckenhorn WU et al (2018) Interrelations of global macroecological patterns in wing and thorax size, sexual size dimorphism, and range size of the Drosophilidae. Ecography 41:1707\u20131717. https:\/\/doi.org\/10.1111\/ecog.03382","journal-title":"Ecography"},{"key":"3739_CR93","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1002\/evl3.153","volume":"4","author":"WG Rostant","year":"2020","unstructured":"Rostant WG, Mason JS, de Coriolis J-C, Chapman T (2020) Resource-dependent evolution of female resistance responses to sexual conflict. Evol Lett 4:54\u201364. https:\/\/doi.org\/10.1002\/evl3.153","journal-title":"Evol Lett"},{"key":"3739_CR94","doi-asserted-by":"publisher","first-page":"1415","DOI":"10.1098\/rspb.1996.0207","volume":"263","author":"L Rowe","year":"1996","unstructured":"Rowe L, Houle D (1996) The lek paradox and the capture of genetic variance by condition dependent traits. Proc R Soc Lond B Biol Sci 263:1415\u20131421. https:\/\/doi.org\/10.1098\/rspb.1996.0207","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"3739_CR95","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1093\/beheco\/ari013","volume":"16","author":"MA Sch\u00e4fer","year":"2005","unstructured":"Sch\u00e4fer MA, Uhl G (2005) Sequential mate encounters: female but not male body size influences female remating behavior. Behav Ecol 16:461\u2013466. https:\/\/doi.org\/10.1093\/beheco\/ari013","journal-title":"Behav Ecol"},{"key":"3739_CR96","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"J Schindelin","year":"2012","unstructured":"Schindelin J, Arganda-Carreras I, Frise E et al (2012) Fiji: an open-source platform for biological-image analysis. Nat Methods 9:676\u2013682. https:\/\/doi.org\/10.1038\/nmeth.2019","journal-title":"Nat Methods"},{"key":"3739_CR97","doi-asserted-by":"publisher","first-page":"1823","DOI":"10.1098\/rspb.2004.2805","volume":"271","author":"N Seddon","year":"2004","unstructured":"Seddon N, Amos W, Mulder RA, Tobias JA (2004) Male heterozygosity predicts territory size, song structure and reproductive success in a cooperatively breeding bird. Proc R Soc Lond B Biol Sci 271:1823\u20131829. https:\/\/doi.org\/10.1098\/rspb.2004.2805","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"3739_CR98","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1007\/s10164-014-0399-z","volume":"32","author":"S Setoguchi","year":"2014","unstructured":"Setoguchi S, Takamori H, Aotsuka T et al (2014) Sexual dimorphism and courtship behavior in Drosophila prolongata. J Ethol 32:91\u2013102. https:\/\/doi.org\/10.1007\/s10164-014-0399-z","journal-title":"J Ethol"},{"key":"3739_CR99","doi-asserted-by":"publisher","DOI":"10.1515\/9780691206882","volume-title":"Mating systems and strategies","author":"SM Shuster","year":"2003","unstructured":"Shuster SM, Wade MJ (2003) Mating systems and strategies. Princeton University Press, Princeton"},{"key":"3739_CR100","doi-asserted-by":"crossref","unstructured":"Thornhill R, Alcock J (1983) The evolution of insect mating systems. Harvard University Press","DOI":"10.4159\/harvard.9780674433960"},{"key":"3739_CR101","doi-asserted-by":"publisher","unstructured":"Tingley D, Yamamoto T, Hirose K et al (2014) mediation: R Package for Causal Mediation Analysis. J Stat Softw 59. https:\/\/doi.org\/10.18637\/jss.v059.i05","DOI":"10.18637\/jss.v059.i05"},{"key":"3739_CR102","doi-asserted-by":"publisher","first-page":"999","DOI":"10.1093\/beheco\/arac067","volume":"33","author":"T Tregenza","year":"2022","unstructured":"Tregenza T, Niemel\u00e4 PT, Rodr\u00edguez-Mu\u00f1oz R, Hopwood PE (2022) Environment and mate attractiveness in a wild insect. Behav Ecol 33:999\u20131006. https:\/\/doi.org\/10.1093\/beheco\/arac067","journal-title":"Behav Ecol"},{"key":"3739_CR103","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/s10905-012-9342-8","volume":"26","author":"E Turiegano","year":"2013","unstructured":"Turiegano E, Monedero I, Pita M et al (2013) Effect of Drosophila melanogaster female size on male mating success. J Insect Behav 26:89\u2013100. https:\/\/doi.org\/10.1007\/s10905-012-9342-8","journal-title":"J Insect Behav"},{"key":"3739_CR104","doi-asserted-by":"publisher","first-page":"481","DOI":"10.1086\/512046","volume":"169","author":"C Vanp\u00e9","year":"2007","unstructured":"Vanp\u00e9 C, Gaillard J, Kjellander P et al (2007) Antler size provides an honest signal of male phenotypic quality in roe deer. Am Nat 169:481\u2013493. https:\/\/doi.org\/10.1086\/512046","journal-title":"Am Nat"},{"key":"3739_CR105","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1145\/2786984.2786995","volume":"19","author":"G Varoquaux","year":"2015","unstructured":"Varoquaux G, Buitinck L, Louppe G et al (2015) Scikit-learn: machine learning without learning the machinery. GetMobile Mob Comp Comm 19:29\u201333. https:\/\/doi.org\/10.1145\/2786984.2786995","journal-title":"GetMobile Mob Comp Comm"},{"key":"3739_CR106","doi-asserted-by":"publisher","first-page":"1145","DOI":"10.1163\/000579510X510593","volume":"147","author":"JL Verdolin","year":"2010","unstructured":"Verdolin JL, Slobodchikoff CN (2010) Male territoriality in a social sciurid, Cynomys gunnisoni: what do patterns of paternity tell us? Behaviour 147:1145\u20131167. https:\/\/doi.org\/10.1163\/000579510X510593","journal-title":"Behaviour"},{"key":"3739_CR107","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1093\/jeb\/voae138","volume":"38","author":"T Verma","year":"2024","unstructured":"Verma T, Das S, Dhodi Lobo S et al (2024) Evolution of mate harm resistance in females from Drosophila melanogaster populations selected for faster development and early reproduction. J Evol Biol 38:111\u2013121. https:\/\/doi.org\/10.1093\/jeb\/voae138","journal-title":"J Evol Biol"},{"key":"3739_CR108","doi-asserted-by":"publisher","first-page":"143","DOI":"10.2307\/1935753","volume":"47","author":"J Verner","year":"1966","unstructured":"Verner J, Willson MF (1966) The influence of habitats on mating systems of North American passerine birds. Ecology 47:143\u2013147. https:\/\/doi.org\/10.2307\/1935753","journal-title":"Ecology"},{"key":"3739_CR109","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1016\/j.anbehav.2016.01.006","volume":"113","author":"C Von Kuerthy","year":"2016","unstructured":"Von Kuerthy C, Taborsky M (2016) Contest versus scramble competition among males pursuing fixed or plastic alternative reproductive tactics. Anim Behav 113:203\u2013212. https:\/\/doi.org\/10.1016\/j.anbehav.2016.01.006","journal-title":"Anim Behav"},{"key":"3739_CR110","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1111\/evo.12580","volume":"69","author":"AJ White","year":"2015","unstructured":"White AJ, Rundle HD (2015) Territory defense as a condition-dependent component of male reproductive success in Drosophila serrata. Evolution 69:407\u2013418. https:\/\/doi.org\/10.1111\/evo.12580","journal-title":"Evolution"},{"key":"3739_CR111","doi-asserted-by":"crossref","unstructured":"Wickham H (2016) ggplot2: elegant graphics for data analysis, Second edition. Springer, Switzerland","DOI":"10.1007\/978-3-319-24277-4_9"},{"key":"3739_CR112","doi-asserted-by":"publisher","first-page":"410","DOI":"10.1111\/1365-2435.12515","volume":"30","author":"S Wigby","year":"2016","unstructured":"Wigby S, Perry JC, Kim Y, Sirot LK (2016) Developmental environment mediates male seminal protein investment in D rosophila melanogaster. Funct Ecol 30:410\u2013419. https:\/\/doi.org\/10.1111\/1365-2435.12515","journal-title":"Funct Ecol"},{"key":"3739_CR113","doi-asserted-by":"publisher","first-page":"1371","DOI":"10.1111\/j.1558-5646.1996.tb03911.x","volume":"50","author":"RH Wiley","year":"1996","unstructured":"Wiley RH, Poston J (1996) Indirect mate choice, competition for mates, and coevolution of the sexes. Evolution 50:1371\u20131381. https:\/\/doi.org\/10.1111\/j.1558-5646.1996.tb03911.x","journal-title":"Evolution"},{"key":"3739_CR114","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1007\/s00265-008-0689-y","volume":"63","author":"BBM Wong","year":"2009","unstructured":"Wong BBM, Svensson PA (2009) Strategic male signalling effort in a desert-dwelling fish. Behav Ecol Sociobiol 63:543\u2013549. https:\/\/doi.org\/10.1007\/s00265-008-0689-y","journal-title":"Behav Ecol Sociobiol"},{"key":"3739_CR115","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/j.anbehav.2014.02.024","volume":"91","author":"JR York","year":"2014","unstructured":"York JR, Baird TA, Haynie ML (2014) Unexpected high fitness payoff of subordinate social tactics in male collared lizards. Anim Behav 91:17\u201325. https:\/\/doi.org\/10.1016\/j.anbehav.2014.02.024","journal-title":"Anim Behav"},{"key":"3739_CR116","doi-asserted-by":"publisher","first-page":"17519","DOI":"10.1073\/pnas.0608211103","volume":"103","author":"A Yurkovic","year":"2006","unstructured":"Yurkovic A, Wang O, Basu AC, Kravitz EA (2006) Learning and memory associated with aggression in Drosophila melanogaster. Proc Natl Acad Sci 103:17519\u201317524. https:\/\/doi.org\/10.1073\/pnas.0608211103","journal-title":"Proc Natl Acad Sci"},{"key":"3739_CR117","doi-asserted-by":"publisher","first-page":"1252","DOI":"10.1111\/jfb.15524","volume":"103","author":"S Zarini","year":"2023","unstructured":"Zarini S, Poli F, Balshine S (2023) Alternative reproductive tactics in goby fishes of the Caspian Sea. J Fish Biol 103:1252\u20131263. https:\/\/doi.org\/10.1111\/jfb.15524","journal-title":"J Fish Biol"},{"key":"3739_CR118","doi-asserted-by":"publisher","first-page":"35","DOI":"10.4161\/fly.19249","volume":"6","author":"L Zwarts","year":"2012","unstructured":"Zwarts L, Versteven M, Callaerts P (2012) Genetics and neurobiology of aggression in Drosophila. Fly (Austin) 6:35\u201348. https:\/\/doi.org\/10.4161\/fly.19249","journal-title":"Fly (Austin)"}],"container-title":["Behavioral Ecology and Sociobiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00265-026-03739-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00265-026-03739-z","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00265-026-03739-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T22:02:46Z","timestamp":1791928966000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00265-026-03739-z"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5]]},"references-count":118,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2026,5]]}},"alternative-id":["3739"],"URL":"https:\/\/doi.org\/10.1007\/s00265-026-03739-z","relation":{},"ISSN":["0340-5443","1432-0762"],"issn-type":[{"value":"0340-5443","type":"print"},{"value":"1432-0762","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5]]},"assertion":[{"value":"28 June 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"31 January 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 April 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 May 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 October 2026","order":6,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":7,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This article was originally published under the subscription model but it is now published under an Open Access license.","order":8,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"61"}}