Abstract
Urbanisation, one of the main driving forces of the decline in arthropod diversity, is a global environmental problem. Urbanisation causes changes to the size, connectivity, structure, and environmental parameters of their natural habitat. Due to a host of novel conditions and situations, high exploratory and risk-taking behaviours are beneficial traits to cope with urban environments. Therefore, we hypothesised that urban spiders should display more exploratory and risk-taking behaviour than their rural conspecifics. We tested 253 individuals of a widespread, forest-associated ground-dwelling wolf spider species, Pardosa alacris, sampled from rural and urban forest sites during their peak activity period, for their locomotory activity, exploratory and risk-taking behaviour by six frequently used behavioural measures. Combining the studied behavioural measures into composite scores using redundancy analysis, we identified two composite variables, the activity-exploration-boldness and the risk-taking behavioural ones. Behaviour measured by the composite activity-exploration-boldness score was significantly repeatable, but not the composite risk-taking behavioural one. There were no urbanisation-related differences in the composite behavioural scores, suggesting that higher exploratory or risk-taking behaviour may not yield fitness benefits in this generalist predator. We found, however, significant sex-specific differences in the composite activity-exploration-boldness behavioural scores. The higher activity, exploratory and boldness in males than females may be explained by their different life-history strategies and sex-specific selective pressures.
Similar content being viewed by others
Data availability
Data used for analyses are available in the Mendeley repository (doi: 10.17632/j6x8hyxgj9.2; https://data.mendeley.com/datasets/j6x8hyxgj9/2).
References
Elmqvist, T. et al. Urbanization in and for the Anthropocene. npj Urban Sustain. 1, 6 (2021).
Hahs, A. K. et al. Urbanisation generates multiple trait syndromes for terrestrial animal taxa worldwide. Nat. Commun. 14, 4751 (2023).
Lowry, H., Lill, A. & Wong, B. B. M. Behavioural responses of wildlife to urban environments. Biol. Rev. 88, 537–549 (2013).
Dall, S. R. X., Houston, A. I. & McNamara, J. M. The behavioural ecology of personality: Consistent individual differences from an adaptive perspective. Ecol. Lett. 7, 734–739 (2004).
Sih, A., Bell, A. & Johnson, J. C. Behavioral syndromes: An ecological and evolutionary overview. Trends Ecol. Evol. 19, 372–378 (2004).
Rodríguez-Muñoz, R., Bretman, A. & Tregenza, T. Guarding males protect females from predation in a wild insect. Curr. Biol. 21, 1716–1719 (2011).
Chown, S. L. & Nicolson, S. Insect Physiological Ecology: Mechanisms and Patterns (Oxford University Press, 2004). https://doi.org/10.1093/acprof:oso/9780198515494.001.0001.
Tew, N. & Hesselberg, T. The effect of wind exposure on the web characteristics of a tetragnathid orb spider. J. Insect Behav. 30, 273–286 (2017).
Kralj-Fišer, S., Hebets, E. A. & Kuntner, M. Different patterns of behavioral variation across and within species of spiders with differing degrees of urbanization. Behav. Ecol. Sociobiol. 71, 125 (2017).
Eötvös, C. B., Magura, T. & Lövei, G. L. A meta-analysis indicates reduced predation pressure with increasing urbanization. Landsc. Urban Plan. 180, 54–59 (2018).
McKinney, M. L. Effects of urbanization on species richness: A review of plants and animals. Urban Ecosyst. 11, 161–176 (2008).
Hedrick, A. V. & Kortet, R. Hiding behaviour in two cricket populations that differ in predation pressure. Anim. Behav. 72, 1111–1118 (2006).
Davis, A. K., Stewart, K., Phelan, C. & Schultz, A. How urban-tolerant are they? Testing prey-capture behavior of introduced Jorō spiders (Trichonephila clavata) next to busy roads. Arthropoda 2, 55–65 (2024).
Schuett, W. et al. Ground beetles in city forests: does urbanization predict a personality trait?. PeerJ 6, e4360 (2018).
Schuett, W., Tregenza, T. & Dall, S. R. X. Sexual selection and animal personality. Biol. Rev. 85, 217–246 (2010).
Kralj-Fišer, S. & Schuett, W. Studying personality variation in invertebrates: Why bother?. Anim. Behav. 91, 41–52 (2014).
Bogyó, D., Magura, T., Simon, E. & Tóthmérész, B. Millipede (Diplopoda) assemblages alter drastically by urbanisation. Landsc. Urban Plan. 133, 118–126 (2015).
Samu, F., Szirányi, A. & Kiss, B. Foraging in agricultural fields: Local ‘sit-and-move’ strategy scales up to risk-averse habitat use in a wolf spider. Anim. Behav. 66, 939–947 (2003).
Michalko, R., Košulič, O., Hula, V. & Surovcová, K. Niche differentiation of two sibling wolf spider species, Pardosa lugubris and Pardosa alacris, along a canopy openness gradient. J. Arachnol 44, 46–51 (2016).
Ingle, K. et al. The effects of overwintering and habitat type on body condition and locomotion of the wolf spider Pardosa alacris. Acta Oecol. 89, 38–42 (2018).
Eliašová, M., Gajdoš, P., Kollár, J., Zuzulová, V. & Šiška, B. Seasonal activity of two ground dwelling arthropod groups at forest stands under different management regimes at the locality Báb (SW Slovakia): the analysis of preliminary results and some suggestion for long-term ecological research. in Towards Climatic Services (eds. Šiška, B., Nejedlík, P. & Eliašová, M.) 26–31 (Slovak University of Agriculture, 2015).
Horváth, R., Magura, T. & Tóthmérész, B. Ignoring ecological demands masks the real effect of urbanization: A case study of ground-dwelling spiders along a rural-urban gradient in a lowland forest in Hungary. Ecol. Res. 27, 1069–1077 (2012).
Magura, T., Horváth, R. & Tóthmérész, B. Effects of urbanization on ground-dwelling spiders in forest patches, in Hungary. Landsc. Ecol. (2010). https://doi.org/10.1007/s10980-009-9445-6.
Magura, T. et al. Urban individuals of three rove beetle species are not more exploratory or risk-taking than rural conspecifics. Insects 13, 757 (2022).
Kortet, R. & Hedrick, A. N. N. A behavioural syndrome in the field cricket Gryllus integer: Intrasexual aggression is correlated with activity in a novel environment. Biol. J. Linn. Soc. 91, 475–482 (2007).
Labaude, S., O’Donnell, N. & Griffin, C. T. Description of a personality syndrome in a common and invasive ground beetle (Coleoptera: Carabidae). Sci. Rep. 8, 17479 (2018).
Magura, T. et al. Are there personality differences between rural vs. urban-living individuals of a specialist ground beetle. Carabus convexus? Insects. 12, 646 (2021).
Tremmel, M. & Müller, C. Insect personality depends on environmental conditions. Behav. Ecol. 24, 386–392 (2013).
Gyuris, E., Feró, O., Tartally, A. & Barta, Z. Individual behaviour in firebugs (Pyrrhocoris apterus). Proc. R. Soc. B Biol. Sci. 278, 628–633 (2011).
Wexler, Y., Subach, A., Pruitt, J. N. & Scharf, I. Behavioral repeatability of flour beetles before and after metamorphosis and throughout aging. Behav. Ecol. Sociobiol. 70, 745–753 (2016).
Bell, A. M., Hankison, S. J. & Laskowski, K. L. The repeatability of behaviour: A meta-analysis. Anim. Behav. 77, 771–783 (2009).
R Core Team. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. (2025).
Dingemanse, N. J. et al. Behavioural syndromes differ predictably between 12 populations of three-spined stickleback. J. Anim. Ecol. 76, 1128–1138 (2007).
Oksanen, J. et al. vegan: Community ecology package. https://doi.org/10.32614/CRAN.package.vegan (2022).
Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Fox, J. & Weisberg, S. An R Companion to Applied Regression (SAGE, 2018).
Venables, W. & Ripley, B. Modern Applied Statistics with S (Springer, 2002).
Zuur, A., Ieno, E. N., Walker, N., Saveliev, A. A. & Smith, G. M. Mixed Effects Models and Extensions in Ecology with R (Springer, 2009). https://doi.org/10.1007/978-0-387-87458-6.
Bartoń, K. & MuMIn Multi-Model Inference. https://doi.org/10.32614/CRAN.package.MuMIn (2025).
Signorell, A. et al. DescTools: Tools for Descriptive Statistics. https://doi.org/10.32614/CRAN.package.DescTools (2021).
Stoffel, M. A., Nakagawa, S. & Schielzeth, H. rptR: Repeatability estimation and variance decomposition by generalized linear mixed-effects models. Methods Ecol. Evol. 8, 1639–1644 (2017).
Sih, A., Bell, A. M., Johnson, J. C. & Ziemba, R. E. Behavioral syndromes: An integrative overview. Q. Rev. Biol. 79, 241–277 (2004).
Budaev, S. V. Using principal components and factor analysis in animal behaviour research: Caveats and guidelines. Ethology 116, 472–480 (2010).
Martinig, A. R. et al. Animal personality: A comparison of standardized assays and focal observations in North American red squirrels. Anim. Behav. 190, 221–232 (2022).
Carere, C. & Maestripieri, D. Animal Personalities: Who Cares and Why? in Animal Personalities - Behavior, Physiology, and Evolution (eds Carere, C. & Maestripieri, D.) 1–10 https://doi.org/10.7208/9780226922065-002 (University of Chicago Press, 2013).
Trabalon, M. Effects of wolf spiders’ captive environment on their locomotor and exploratory behaviours. Insects 13, 135 (2022).
Liedtke, J., Redekop, D., Schneider, J. M. & Schuett, W. Early environmental conditions shape personality types in a jumping spider. Front. Ecol. Evol. 3, 134 (2015).
Kralj-Fišer, S. & Schneider, J. M. Individual behavioural consistency and plasticity in an urban spider. Anim. Behav. 84, 197–204 (2012).
Bengston, S. E. & Dornhaus, A. Be meek or be bold? A colony-level behavioural syndrome in ants. Proc. R. Soc. B Biol. Sci. 281, 20140518 (2014).
Jones, C. & DiRienzo, N. Behavioral variation post-invasion: Resemblance in some, but not all, behavioral patterns among invasive and native praying mantids. Behav. Processes. 153, 92–99 (2018).
Louca, V., Savorelli, S., Vuorio, N. & Fisher, D. N. Cockroach social network position does not influence exploration tendency. J. Ethol. 44, 75–86 (2026).
Andrew, R. J. Recognition processes and behavior, with special reference to effects of testosterone on persistence. Adv. Study Behav. 4, 175–208 (1972).
Šramel, N., Kablar, D., Debes, P. V. & Kralj-Fišer, S. Sex differences in the behavioural traits across ontogenetic stages in a sexually size dimorphic spider. Anim. Behav. 207, 183–189 (2024).
Cordellier, M., Schneider, J. M., Uhl, G. & Posnien, N. Sex differences in spiders: From phenotype to genomics. Dev. Genes Evol. 230, 155–172 (2020).
Wu, L., Zhang, H., He, T., Liu, Z. & Peng, Y. Factors influencing sexual cannibalism and its benefit to fecundity and offspring survival in the wolf spider Pardosa pseudoannulata (Araneae: Lycosidae). Behav. Ecol. Sociobiol. 67, 205–212 (2013).
Yli-Renko, M., Pettay, J. E. & Vesakoski, O. Sex and size matters: Selection on personality in natural prey-predator interactions. Behav. Processes. 148, 20–26 (2018).
Persons, M. H. Hunger effects on foraging responses to perceptual cues in immature and adult wolf spiders (Lycosidae). Anim. Behav. 57, 81–88 (1999).
Mezőfi, L., Markó, V., Taranyi, D. Á. & Markó, G. Sex-specific life-history strategies among immature jumping spiders: Differences in body parameters and behavior. Curr. Zool. 69, 535–551 (2023).
Kralj-Fišer, S., Schneider, J. M., Kuntner, M., Laskowski, K. & Garcia-Gonzalez, F. The genetic architecture of behavioral traits in a spider. Ecol. Evol. 11, 5381–5392 (2021).
Audet, J.-N., Ducatez, S. & Lefebvre, L. The town bird and the country bird: Problem solving and immunocompetence vary with urbanization. Behav. Ecol. 27, 637–644 (2015).
Caizergues, A. E., Grégoire, A., Choquet, R., Perret, S. & Charmantier, A. Are behaviour and stress-related phenotypes in urban birds adaptive? J. Anim. Ecol. 91, 1627–1641 (2022).
Vines, A. & Lill, A. Boldness and urban dwelling in little ravens. Wildl. Res. 42, 590–597 (2016).
Dammhahn, M., Mazza, V., Schirmer, A., Göttsche, C. & Eccard, J. A. Of city and village mice: behavioural adjustments of striped field mice to urban environments. Sci. Rep. 10, 13056 (2020).
Mazza, V., Dammhahn, M., Lösche, E. & Eccard, J. A. Small mammals in the big city: Behavioural adjustments of non-commensal rodents to urban environments. Glob. Chang. Biol. 26, 6326–6337 (2020).
von Merten, S. et al. Urban populations of shrews show larger behavioural differences among individuals than rural populations. Anim. Behav. 187, 35–46 (2022).
Rech, F., Narimanov, N., Bauer, T. & Schirmel, J. Urbanization increases fluctuating asymmetry and affects behavioral traits of a common grasshopper. Ecol. Evol. 12, e9658 (2022).
Jacquier, L., Molet, M. & Doums, C. Urban colonies are less aggressive but forage more than their forest counterparts in the ant Temnothorax nylanderi. Anim. Behav. 199, 11–21 (2023).
Trigos-Peral, G. et al. Urban abiotic stressors drive changes in the foraging activity and colony growth of the black garden ant Lasius niger. Sci. Total Environ. 915, 170157 (2024).
Halpin, R. N. & Johnson, J. C. A continuum of behavioral plasticity in urban and desert black widows. Ethology 120, 1237–1247 (2014).
Réale, D., Reader, S. M., Sol, D., McDougall, P. T. & Dingemanse, N. J. Integrating animal temperament within ecology and evolution. Biol. Rev. 82, 291–318 (2007).
Kaiser, A., Merckx, T. & Van Dyck, H. An experimental test of changed personality in butterflies from anthropogenic landscapes. Behav. Ecol. Sociobiol. 74, 86 (2020).
Markowski, M. et al. Genetic structure of urban and non-urban populations differs between two common parid species. Sci. Rep. 11, 10428 (2021).
Sappington, T. W. Aseasonal, undirected migration in insects: ‘Invisible’ but common. iScience 27, 110040 (2024).
Bengston, S. E., Pruitt, J. N. & Riechert, S. E. Differences in environmental enrichment generate contrasting behavioural syndromes in a basal spider lineage. Anim. Behav. 93, 105–110 (2014).
Mitchell, D. J. & Houslay, T. M. Context-dependent trait covariances: How plasticity shapes behavioral syndromes. Behav. Ecol. 32, 25–29 (2020).
Wong, B. B. M. & Candolin, U. Behavioral responses to changing environments. Behav. Ecol. 26, 665–673 (2014).
Wolf, M. & Weissing, F. J. An explanatory framework for adaptive personality differences. Philos. Trans. R. Soc. Lond. B Biol. Sci. 365, 3959–3968 (2010).
Bergmüller, R. & Taborsky, M. Animal personality due to social niche specialisation. Trends Ecol. Evol. 25, 504–511 (2010).
Acknowledgements
We thank Réka Csicsek and Dávid D. Nagy for your help during the study, as well as the Department of Green Infrastructure of the Mayor’s Office of Debrecen, especially Orsolya Hamecz, for the permission to conduct the study. This research was funded by the Hungarian National Research, Development and Innovation Fund (grant number OTKA K-146628). Authorship is by the “first-and-last-author-emphasis” (FLAE) principle.
Funding
Open access funding provided by University of Debrecen.
Author information
Authors and Affiliations
Contributions
T.M.: conceptualisation, data curation, formal analysis, funding acquisition, investigation, methodology, supervision, project administration, resources, validation, visualisation, writing—original draft, writing—review and editing. R.H.: conceptualisation, investigation, methodology, writing—original draft, writing—review and editing. S.M.: investigation, methodology, writing—review and editing. M.T.: investigation, methodology, writing—review and editing. S.F.K.: methodology, visualisation, writing—review and editing. G.L.L.: conceptualisation, methodology, supervision, validation, writing—original draft, writing—review and editing.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Magura, T., Horváth, R., Mizser, S. et al. Sex-specific but not urbanisation-related behavioural differences in a wolf spider, Pardosa alacris. Sci Rep (2026). https://doi.org/10.1038/s41598-026-41239-2
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-026-41239-2


