Abstract
Although conflict is often adaptive and necessary to secure limited resources, it is also frequently a costly endeavor. Signals that reliably communicate competitive ability are commonly employed by animals to reduce conflict costs. Both male and female signals have the capacity to serve as honest indicators of competitive ability, but the extent to which this occurs in females has received relatively limited attention. We studied how natural feather coloration and experimentally manipulated plumage ornamentation in female white-shouldered fairywrens (Malurus alboscapulatus moretoni) relates to their aggressive behavior. These fairywrens are a useful study system in that female, but not male coloration varies throughout New Guinea. We tested behavioral responses to simulated rivals both prior to and after plumage manipulation via two distinct behavioral assays: simulated territorial intrusions (where both sexes jointly respond to rival intruders) and mirror image simulation (where females are isolated from their mate). Plumage manipulation treatments had no measurable impact on female aggression during mirror image simulation tests, though aggression did decrease over the course of multiple assays. Similarly, using simulated territorial intrusion assays, we found no difference in female aggression with respect to both natural coloration and manipulation treatment. Finally, our correlative analysis of natural feather color also revealed no detectable relationship. These findings suggest that female white scapular coloration is unlikely to function as a signal of competitive ability in this tropical species with derived female ornamentation and we discuss possible alternative explanations.
Significance statement
Like their showy male counterparts, female animals may use plumage-based signals to mediate aggressive encounters. Determining if and how signals used in female-female competition may mitigate the costs of agonistic encounters is particularly important, as it is thought to be a key driver of female signal evolution. We used a repeated-measures design and two distinct behavioral assays to explore how both manipulated and natural feather coloration relate to aggression in female white-shouldered fairywrens of Papua New Guinea – a species with female, but not male, plumage polymorphisms. We were unable to detect any differences in aggression with respect to female plumage coloration despite two distinct behavioral assays that mimic distinct social contexts, suggesting that female plumage coloration may function in other contexts or lack a modern signaling function altogether in this system.
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Data availability
Data and associated R scripts are available via Dryad: DOI:10.5061/dryad.6djh9w12k.
References
Amundsen T, Forsgren E (2001) Male mate choice selects for female coloration in a fish. P Natl Acad Sci USA 98:13155–13160. https://doi.org/10.1073/pnas.211439298
Audacity Team (2018) Audacity(R), v 2.2.2, https://audacityteam.org/
Autumn K, Ryan MJ, Wake DB (2002) Integrating historical and mechanistic biology enhances the study of adaptation. Q Rev Biol 77:383–408. https://doi.org/10.1086/344413
Bates D, Mächler M, Bolker BM, Walker SC (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1–48. https://doi.org/10.18637/jss.v067.i01
Beck ML, Hopkins WA (2019) The relationship between plumage coloration and aggression in female tree swallows. J Avian Biol 50:e01910. https://doi.org/10.1111/jav.01910
Berzins LL, Dawson RD (2018) Experimentally altered plumage brightness of female tree swallows (Tachycineta bicolor) influences nest site retention and reproductive success. Can J Zool 96:600–607. https://doi.org/10.1139/cjz-2017-0156
Biernaskie JM, Grafen A, Perry JC (2014) The evolution of index signals to avoid the cost of dishonesty. Proc R Soc B 281:20140876. https://doi.org/10.1098/rspb.2014.0876
Boersma J, Enbody ED, Jones JA, Nason D, Lopez-Contreras KJ, Schwabl H (2020) Testosterone induces plumage ornamentation followed by enhanced territoriality in a female songbird. Behav Ecol 31:1233–1241. https://doi.org/10.1093/beheco/araa077
Bonduriansky R (2001) The evolution of male mate choice in insects: a synthesis of ideas and evidence. Biol Rev 76:305–339. https://doi.org/10.1017/S1464793101005693
Cain KE, Rosvall KA (2014) Next steps for understanding the selective relevance of female-female competition. Front Ecol Evol 2:32. https://doi.org/10.3389/fevo.2014.00032
Cardoso GC, Mota PG (2010) Evolution of female carotenoid coloration by sexual constraint in Carduelis finches. BMC Evol Biol 10:82. https://doi.org/10.1186/1471-2148-10-82
Cassey P, Ewen JG, Blackburn TM, Hauber ME, Voryobyev M, Marshall NJ (2008) Eggshell colour does not predict measures of maternal investment in eggs of Turdus thrushes. Naturwissenschaften 95:713–721. https://doi.org/10.1007/s00114-008-0376-x
Chaine AS, Shizuka D, Block TA, Zhang L, Lyon BE (2018) Manipulating badges of status only fools strangers. Ecol Lett 21:1477–1485. https://doi.org/10.1111/ele.13128
Clutton-Brock T (2007) Sexual selection in males and females. Science 318:1882–1885. https://doi.org/10.1126/science.1133311
Cockrem JF, Silverin B (2002) Variation within and between birds in corticosterone responses of great tits (Parus major). Gen Comp Endocrinol 125:197–206. https://doi.org/10.1006/gcen.2001.7750
Cole GL, Endler JA (2016) Male courtship decisions are influenced by light environment and female receptivity. Proc R Soc B 283:20160861. https://doi.org/10.1098/rspb.2016.0861
Courtiol A, Etienne L, Feron R, Godelle B, Rousset F (2016) The evolution of mutual mate choice under direct benefits. Am Nat 188:521–538. https://doi.org/10.1086/688658
Cuthill IC (2006) Color perception. In: Hill GE, McGraw KJ (eds) Bird Coloration I: Mechanisms and Measurements. Harvard University Press, Cambridge, MA, pp 3–40
Dale J, Dey C, Delhey K, Kempenaers B, Valcu M (2015) The effects of life-history and social selection on male and female plumage coloration. Nature 527:367–370. https://doi.org/10.1038/nature15509
Delhey K, Delhey V, Kempenaers B, Peters A (2015) A practical framework to analyze variation in animal colors using visual models. Behav Ecol 26:367–375. https://doi.org/10.1093/beheco/aru198
Delhey K, Hall M, Kingma SA, Peters A (2013) Increased conspicuousness can explain the match between visual sensitivities and blue plumage colours in fairy-wrens. Proc R Soc B 280:20121771. https://doi.org/10.1098/rspb.2012.1771
Delhey K, Peters A (2008) Quantifying variability of avian colours: are signalling traits more variable? PLoS ONE 3:e1689. https://doi.org/10.1371/journal.pone.0001689
Diamant ES, Falk JJ, Rubenstein DR (2021) Male-like female morphs in hummingbirds: the evolution of a widespread sex-limited plumage polymorphism. Proc R Soc B 288:20203004. https://doi.org/10.1098/rspb.2020.3004
Doutrelant C, Fargevieille A, Grégoire A (2020) Evolution of female coloration: what have we learned from birds in general and blue tits in particular. Adv Stud Behav 52:123–202. https://doi.org/10.1016/bs.asb.2020.03.001
Driskell AC, Norman JA, Pruett-Jones S, Mangall E, Sonsthagen S, Christidis L (2011) A multigene phylogeny examining evolutionary and ecological relationships in the Australo-Papuan wrens of the subfamily Malurinae (Aves). Mol Phylogenet Evol 60:480–485. https://doi.org/10.1016/j.ympev.2011.03.030
Enbody ED, Boersma J, Jones JA et al (2019) Social organisation and breeding biology of the white-shouldered fairywren (Malurus alboscapulatus). Emu 119:274–285. https://doi.org/10.1080/01584197.2019.1595663
Enbody ED, Boersma J, Schwabl H, Karubian J (2018) Female ornamentation is associated with elevated aggression and testosterone in a tropical songbird. Behav Ecol 29:1056–1066. https://doi.org/10.1093/beheco/ary079
Endler JA, Mielke PW (2005) Comparing entire colour patterns as birds see them. Biol J Linn Soc 86:405–431. https://doi.org/10.1111/j.1095-8312.2005.00540.x
Ferns PN, Hinsley SA (2004) Immaculate tits: head plumage pattern as an indicator of quality in birds. Anim Behav 67:261–272. https://doi.org/10.1016/j.anbehav.2003.05.006
Fitzpatrick CL, Servedio MR (2017) Male mate choice, male quality, and the potential for sexual selection on female traits under polygyny. Evolution 71:174–183. https://doi.org/10.1111/evo.13107
Fitzpatrick CL, Servedio MR (2018) The evolution of male mate choice and female ornamentation: a review of mathematical models. Curr Zool 64:323–333. https://doi.org/10.1093/cz/zoy029
Fitzpatrick S, Berglund A, Rosenqvist G (1995) Ornaments or offspring: costs to reproductive success restrict sexual selection processes. Biol J Linn Soc 55:251–260. https://doi.org/10.1111/j.1095-8312.1995.tb01063.x
Freeman BG, Scholer MN, Boehm MMA, Heavyside J, Schluter D (2020) Adaptation and latitudinal gradients in species interactions: nest predation in birds. Am Nat 196:E160–E166. https://doi.org/10.1086/711415
Gallup GG Jr (1968) Mirror-image stimulation. Psychol Bull 70:782–793. https://doi.org/10.1037/h0026777
Gould SJ, Vrba ES (1982) Exaptation – a missing term in the science of form. Paleobiology 8:4–15. https://doi.org/10.2307/2400563
Griggio M, Valera F, Casas A, Pilastro A (2005) Males prefer ornamented females: a field experiment of male choice in the rock sparrow. Anim Behav 69:1243–1250. https://doi.org/10.1016/j.anbehav.2004.10.004
Hall ML, Peters A (2008) Coordination between the sexes for territorial defence in a duetting fairy-wren. Anim Behav 76:65–73. https://doi.org/10.1016/j.anbehav.2008.01.010
Hart NS (2001) Variations in cone photoreceptor abundance and the visual ecology of birds. J Comp Physiol A 187:685–697. https://doi.org/10.1007/s00359-001-0240-3
Hegyi G, Kötél D, Laczi M (2015) Direct benefits of mate choice: a meta-analysis of plumage colour and offspring feeding rates in birds. Sci Nat 102:62. https://doi.org/10.1007/s00114-015-1311-6
Heinsohn R, Legge S, Endler JA (2005) Evolution: extreme reversed sexual dichromatism in a bird without sex role reversal. Science 309:617–619. https://doi.org/10.1126/science.1112774
Hill GE, McGraw KJ (eds) (2006) Bird Coloration, Vol 1: Mechanisms and Measurements. Harvard University Press, Cambridge, MA
Hofmann CM, Cronin TW, Omland KE (2008) Evolution of sexual dichromatism. 1. Convergent losses of elaborate female coloration in new world orioles (Icterus spp.). Auk 125:790–795. https://doi.org/10.1525/auk.2008.07113
Jawor JM, Gray N, Beall SM, Breitwisch R (2004) Multiple ornaments correlate with aspects of condition and behaviour in female northern cardinals, Cardinalis cardinalis. Anim Behav 67:875–882. https://doi.org/10.1016/j.anbehav.2003.05.015
Johnson AE, Jordan Price J, Pruett-Jones S (2013) Different modes of evolution in males and females generate dichromatism in fairy-wrens (Maluridae). Ecol Evol 3:3030–3046. https://doi.org/10.1002/ece3.686
Johnstone RA, Norris K (1993) Badges of status and the cost of aggression. Behav Ecol Sociobiol 32:127–134
Jones JA, Odom KJ, Hoppe IR, Nason D, Ketaloya S, Karubian J (2021) Correlated evolution of distinct signals associated with increased social selection in female white-shouldered fairywrens. Ecol Evol 11:17352–17363. https://doi.org/10.1002/ece3.8370
Karubian J (2013) Female ornamentation in Malurus fairy-wrens: a hidden evolutionary gem for understanding female perspectives on social and sexual selection. Emu 113:248–258. https://doi.org/10.1071/MU12093
Karubian J, Swaddle JP, Varian-Ramos CW, Webster MS (2009) The relative importance of male tail length and nuptial plumage on social dominance and mate choice in the red-backed fairy-wren Malurus melanocephalus: evidence for the multiple receiver hypothesis. J Avian Biol 40:559–568. https://doi.org/10.1111/j.1600-048X.2009.04572.x
Kelly CD, Godin J-GJ (2001) Predation risk reduces male-male sexual competition in the Trinidadian guppy (Poecilia reticulata). Behav Ecol Sociobiol 51:95–100. https://doi.org/10.1007/s002650100410
Kraaijeveld K (2014) Reversible trait loss: the genetic architecture of female ornaments. Annu Rev Ecol Evol S 45:159–177. https://doi.org/10.1146/annurev-ecolsys-120213-091550
Lande R (1980) Sexual dimorphism, sexual selection, and adaption in polygenic characters. Evolution 34:292–305
LeBas NR (2006) Female finery is not for males. Trends Ecol Evol 21:170–173. https://doi.org/10.1016/j.tree.2006.01.005
LeBas NR, Hockham LR, Ritchie MG (2003) Nonlinear and correlational sexual selection on “honest” female ornamentation. Proc R Soc Lond B 270:2159–2165. https://doi.org/10.1098/rspb.2003.2482
Leitão AV, Hall ML, Delhey K, Mulder RA (2019a) Female and male plumage colour signals aggression in a dichromatic tropical songbird. Anim Behav 150:285–301. https://doi.org/10.1016/j.anbehav.2019.01.025
Leitão AV, Hall ML, Venables B, Mulder RA (2019b) Ecology and breeding biology of a tropical bird, the Lovely Fairy-Wren (Malurus amabilis). Emu 119:1–13. https://doi.org/10.1080/01584197.2018.1498745
Lipshutz SE, Rosvall KA (2021) Nesting strategy shapes territorial aggression but not testosterone: a comparative approach in female and male birds. Horm Behav 133:104995. https://doi.org/10.1016/j.yhbeh.2021.104995
López-Idiáquez D, Vergara P, Fargallo JA, Martínez-Padilla J (2016) Female plumage coloration signals status to conspecifics. Anim Behav 121:101–106. https://doi.org/10.1016/j.anbehav.2016.08.020
Maia R, Gruson H, Endler JA, White TE (2019) pavo 2: New tools for the spectral and spatial analysis of colour in R. Methods Ecol Evol 10:1097–1107. https://doi.org/10.1111/2041-210X.13174
Mason NA, Bowie RCK (2020) Plumage patterns: ecological functions, evolutionary origins, and advances in quantification. Auk 137:ukaa060. doi: https://doi.org/10.1093/auk/ukaa060
Maynard Smith J, Harper DG (1988) The evolution of aggression: can selection generate variability? Phil Trans R Soc B 319:557–570. https://doi.org/10.1098/rstb.1988.0065
Maynard Smith J, Price GR (1973) The logic of animal conflict. Nature 246:15–18. https://doi.org/10.1038/246015a0
Medina I, Delhey K, Peters A, Cain KE, Hall ML, Mulder RA, Langmore NE (2017) Habitat structure is linked to the evolution of plumage colour in female, but not male, fairy-wrens. BMC Evol Biol 17:35. https://doi.org/10.1186/s12862-016-0861-3
Midamegbe A, Grégoire A, Perret P, Doutrelant C (2011) Female-female aggressiveness is influenced by female coloration in blue tits. Anim Behav 82:245–253. https://doi.org/10.1016/j.anbehav.2011.04.020
Montgomerie R (2006) Analyzing colors. In: Hill GE, McGraw KJ (eds) Bird Coloration I: Mechanisms and Measurements. Harvard University Press, Cambridge, MA, pp 90–147
Murphy TG, Rosenthal MF, Montgomerie R, Tarvin KA (2009) Female American goldfinches use carotenoid-based bill coloration to signal status. Behav Ecol 20:1348–1355. https://doi.org/10.1093/beheco/arp140
Nordeide JT, Kekäläinen J, Janhunen M, Kortet R (2013) Female ornaments revisited – are they correlated with offspring quality? J Anim Ecol 82:26–38. https://doi.org/10.1111/1365-2656.12021
Ödeen A, Håstad O (2013) The phylogenetic distribution of ultraviolet sensitivity in birds. BMC Evol Biol 13:36
Ödeen A, Pruett-Jones S, Driskell AC, Armenta JK, Håstad O (2012) Multiple shifts between violet and ultraviolet vision in a family of passerine birds with associated changes in plumage coloration. Proc R Soc Lond B 279:1269–1276. https://doi.org/10.1098/rspb.2011.1777
Odom KJ, Cain KE, Hall ML et al (2021) Sex role similarity and sexual selection predict male and female song elaboration and dimorphism in fairy-wrens. Ecol Evol 11:17901–17919. https://doi.org/10.1002/ece3.8378
Odom KJ, Hall ML, Riebel K, Omland KE, Langmore NE (2014) Female song is widespread and ancestral in songbirds. Nat Commun 5:3379. https://doi.org/10.1038/ncomms4379
Olsson P, Lind O, Kelber A (2018) Chromatic and achromatic vision: parameter choice and limitations for reliable model predictions. Behav Ecol 29:273–282. https://doi.org/10.1093/beheco/arx133
Omland KE, Hofmann CM (2006) Adding color to the past: ancestral-state reconstruction of coloration. In: Hill GE, McGraw KJ (eds) Bird Coloration II: Function and Evolution. Harvard University Press, Cambridge, MA, pp 417–454
Parker GA (1974) Assessment strategy and the evolution of fighting behaviour. J Theor Biol 47:223–243. https://doi.org/10.1016/0022-5193(74)90111-8
R Core Team (2021) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, https://www.R-project.org/.
Revelle W (2021) psych: procedures for psychological, psychometric, and personality research. R package v2.1.6, https://personality-project.org/r/psych/
Riebel K, Odom KJ, Langmore NE, Hall ML (2019) New insights from female bird song: towards an integrated approach to studying Male and female communication roles. Biol Lett 15:20190059. https://doi.org/10.1098/rsbl.2019.0059
Rohwer S (1982) The evolution of reliable and unreliable badges of fighting ability. Am Zool 22:531–546. https://doi.org/10.1093/icb/22.3.531
Rosvall KA (2011) Intrasexual competition in females: evidence for sexual selection? Behav Ecol 22:1131–1140. https://doi.org/10.1093/beheco/arr106
Rowley I, Russell E (1997) Bird families of the world: fairy-wrens and Grasswrens. Oxford University Press, Oxford, UK
Rubenstein DR, Lovette IJ (2009) Reproductive skew and selection on female ornamentation in social species. Nature 462:786–789. https://doi.org/10.1038/nature08614
Santos ESA, Scheck D, Nakagawa S (2011) Dominance and plumage traits: meta-analysis and metaregression analysis. Anim Behav 82:3–19. https://doi.org/10.1016/j.anbehav.2011.03.022
Schneider CA, Rasband WS, Eliceiri KW (2012) NIH image to ImageJ: 25 years of image analysis. Nat Methods 9:671–675. https://doi.org/10.1038/nmeth.2089
Senar JC (2006) Bird colors as intrasexual signals of aggression and dominance. In: Hill GE, McGraw KJ (eds) Bird Coloration II: Function and Evolution. Harvard University Press, Cambridge, MA, USA, pp 125–193
Slater PJB, Mann NI (2004) Why do the females of many bird species sing in the tropics? J Avian Biol 35:289–294. https://doi.org/10.1111/j.0908-8857.2004.03392.x
Stockley P, Bro-Jørgensen J (2011) Female competition and its evolutionary consequences in mammals. Biol Rev 86:341–366. https://doi.org/10.1111/j.1469-185X.2010.00149.x
Stutchbury BJM, Morton ES (2001) Behavioral ecology of tropical birds. Academic Press, London, UK
Stutchbury BJM, Morton ES (2008) Recent advances in the behavioral ecology of tropical birds. Wilson J Ornithol 120:26–37. https://doi.org/10.1676/07-018.1
Swaddle JP, Karubian J, Pruett-jones S (2000) A novel evolutionary pattern of reversed sexual dimorphism in fairy wrens: implications for sexual selection. Behav Ecol 11:345–349. https://doi.org/10.1093/beheco/11.3.345
Taff CC, Zimmer C, Scheck D, Ryan TA, Houtz JL, Smee MR, Hendry TA, Vitousek MN (2021) Plumage manipulation alters associations between behaviour, physiology, the internal microbiome and fitness. Anim Behav 178:11–36. https://doi.org/10.1016/j.anbehav.2021.05.012
Templeton CN, Akçay Ç, Campbell SE, Beecher MD (2012) Soft song is a reliable signal of aggressive intent in song sparrows. Behav Ecol Sociobiol 66:1503–1509. https://doi.org/10.1007/s00265-012-1405-5
Tobias JA, Montgomerie R, Lyon BE (2012) The evolution of female ornaments and weaponry: social selection, sexual selection and ecological competition. Phil Trans R Soc B 367:2274–2293. https://doi.org/10.1098/rstb.2011.0280
Vorobyev M, Osorio D (1998) Receptor noise as a determinant of colour thresholds. Proc R Soc Lond B 265:351–358. https://doi.org/10.1098/rspb.1998.0302
Vorobyev M, Osorio D, Bennett AT, Marshall NJ, Cuthill IC (1998) Tetrachromacy, oil droplets and bird plumage colours. J Comp Physiol A 183:621–633. https://doi.org/10.1007/s003590050286
Weaver RJ, Koch RE, Hill GE (2017) What maintains signal honesty in animal colour displays used in mate choice? Phil Trans R Soc B 372:20160343. https://doi.org/10.1098/rstb.2016.0343
Webb WH, Brunton DH, Aguirre JD, Thomas DB, Valcu M, Dale J (2016) Female song occurs in songbirds with more elaborate female coloration and reduced sexual dichromatism. Front Ecol Evol 4:22. https://doi.org/10.3389/fevo.2016.00022
Weiss SL (2006) Female-specific color is a signal of quality in the striped plateau lizard (Sceloporus virgatus). Behav Ecol 17:726–732. https://doi.org/10.1093/beheco/arl001
West-Eberhard MJ (1979) Sexual selection, social competition, and evolution. P Am Philos Soc 123:222–234
West-Eberhard MJ (1983) Sexual selection, social competition, and speciation. Q Rev Biol 58:155–183
Wiens JJ (2001) Widespread loss of sexually selected traits: how the peacock lost its spots. Trends Ecol Evol 16:517–523. https://doi.org/10.1016/S0169-5347(01)02217-0
Wilkins MR, Odom KJ, Benedict L, Safran RJ (2020) Analysis of female song provides insight into the evolution of sex differences in a widely studied songbird. Anim Behav 168:69–82. https://doi.org/10.1016/j.anbehav.2020.07.018
Acknowledgements
We thank the locals of Podagha Village (Milne Bay Province, Papua New Guinea), without whom none of this research would have been possible. We are grateful for their hospitality and for their logistical and in-field support. We thank the Milne Bay provincial government for permits and permissions and the National Research Institute for their assistance in acquiring country-level research permits (#99902100765) and visas. Finally, we are grateful to Ian Ross Hoppe, Annelise Blanchette, Kevin McGraw, and two anonymous reviewers for their insights and comments on earlier versions of this manuscript.
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This work was supported by the National Science Foundation (IOS-1354133) awarded to JK, the American Ornithological Society to JAJ and JB, the Animal Behaviour Society to JAJ, and the Department of Ecology and Evolutionary Biology of Tulane University to JAJ. Funders have had no influence on the content of the submitted manuscript, nor do they require approval for the final manuscript to be published.
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Our study was carried out in strict accordance with the guidelines established by the Tulane University Institutional Animal Care and Use Committee (#0395R2) as well as in adherence to research permits from the Conservation and Environment Protection Authority of Papua New Guinea (#99902100765). All birds were captured, processed, exposed to one (or two) mirror assays, and then released in under one hour. We continuously monitored our mist-nets and removed birds immediately upon hitting the net. All individuals involved in the removal of birds from mist-nets were trained in the appropriate way to handle birds prior to their handling. All manipulated birds recaptured had marker residue cleaned with rubbing alcohol and feathers plucked to induce healthy molting of a normal shoulder patch. Through routine monitoring of the birds throughout the season, we confirmed that the plumage manipulation did not have any lasting effects (i.e., shoulders were visibly naturally white again a few weeks after treatment).
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Jones, J.A., Boersma, J., Liu, J. et al. Female ornamentation does not predict aggression in a tropical songbird. Behav Ecol Sociobiol 76, 57 (2022). https://doi.org/10.1007/s00265-022-03165-x
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DOI: https://doi.org/10.1007/s00265-022-03165-x