{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T22:39:44Z","timestamp":1790375984220,"version":"4.1.0"},"reference-count":51,"publisher":"MDPI AG","issue":"23","license":[{"start":{"date-parts":[[2025,11,29]],"date-time":"2025-11-29T00:00:00Z","timestamp":1764374400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFC3205903"],"award-info":[{"award-number":["2023YFC3205903"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012428","name":"Central Public-interest Scientific Institution Basal Research Fund, CAFS","doi-asserted-by":"crossref","award":["2023TD09"],"award-info":[{"award-number":["2023TD09"]}],"id":[{"id":"10.13039\/501100012428","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>In freshwater fish, group behavior is ecologically critical for daily activities such as predator avoidance. However, species with varying shoaling preferences exhibit divergent behavioral responses under different environmental conditions. This study investigated the behavioral responses of three shoaling species (Moenkhausia costae, Puntius tetrazona, and Myxocyprinus asiaticus) and three non-shoaling species (Trichogaster trichopterus, Micropterus salmoides, and Cichlasoma managuense) to simulated predation in either an open arena or a six-arm maze with shelter available. Our findings reveal that, in open water, shoaling species employ a dual strategy against predators: maintaining high group cohesion while increasing swimming speed and acceleration. This exploits the confusion effect to mitigate individual predation risk. In contrast, non-shoaling species do not engage in evasive maneuvers; instead, they adopt a cryptic strategy by minimizing activity and often freezing in place to avoid detection. In the six-arm maze, shoaling species consistently employed group coordination strategies, whereas non-shoaling species primarily relied on shelter concealment or reduced activity. Notably, shoaling species maintained high cohesion, synchronization, and activity levels across both open and complex habitats, using coordinated movement to facilitate collective escape. Together, our findings demonstrate that habitat complexity and social tendencies jointly determine how fishes trade off risk and safety. This work provides new insights into the adaptive evolution of social behavior in dynamic aquatic ecosystems.<\/jats:p>","DOI":"10.3390\/ani15233447","type":"journal-article","created":{"date-parts":[[2025,12,3]],"date-time":"2025-12-03T08:44:28Z","timestamp":1764751468000},"page":"3447","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Anti-Predator Strategies in Fish with Contrasting Shoaling Preferences Across Different Contexts"],"prefix":"10.3390","volume":"15","author":[{"given":"Zixi","family":"Lu","sequence":"first","affiliation":[{"name":"Laboratory of Evolutionary Physiology and Behavior, Chongqing Key Laboratory of Conservation and Utilization of Freshwater Fishes, Animal Biology Key Laboratory of Chongqing Education Commission of China, Chongqing Normal University, Chongqing 401331, China"},{"name":"National Agricultural Science Observing and Experimental Station of Chongqing, Fishes Conservation and Utilization in the Upper Reaches of the Yangtze River Key Laboratory of Sichuan Province, Yangtze River Fisheries Research Institute, Chinese Academy of Fishery Science, Wuhan 430223, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-7789-5870","authenticated-orcid":false,"given":"Wuxin","family":"Li","sequence":"additional","affiliation":[{"name":"Laboratory of Evolutionary Physiology and Behavior, Chongqing Key Laboratory of Conservation and Utilization of Freshwater Fishes, Animal Biology Key Laboratory of Chongqing Education Commission of China, Chongqing Normal University, Chongqing 401331, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiuhong","family":"Zhang","sequence":"additional","affiliation":[{"name":"Laboratory of Evolutionary Physiology and Behavior, Chongqing Key Laboratory of Conservation and Utilization of Freshwater Fishes, Animal Biology Key Laboratory of Chongqing Education Commission of China, Chongqing Normal University, Chongqing 401331, China"},{"name":"Sichuan Zigong Ecological Environment Monitoring Center, Zigong 643000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinbin","family":"Duan","sequence":"additional","affiliation":[{"name":"National Agricultural Science Observing and Experimental Station of Chongqing, Fishes Conservation and Utilization in the Upper Reaches of the Yangtze River Key Laboratory of Sichuan Province, Yangtze River Fisheries Research Institute, Chinese Academy of Fishery Science, Wuhan 430223, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7665-9037","authenticated-orcid":false,"given":"Shijian","family":"Fu","sequence":"additional","affiliation":[{"name":"Laboratory of Evolutionary Physiology and Behavior, Chongqing Key Laboratory of Conservation and Utilization of Freshwater Fishes, Animal Biology Key Laboratory of Chongqing Education Commission of China, Chongqing Normal University, Chongqing 401331, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,11,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.1098\/rstb.2007.2103","article-title":"Hypoxia and the Antipredator Behaviours of Fishes","volume":"362","author":"Domenici","year":"2007","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S0149-7634(01)00042-2","article-title":"Antipredator Responses and Defensive Behavior: Ecological and Ethological Approaches for the Neurosciences","volume":"25","author":"Kavaliers","year":"2001","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1093\/beheco\/arm109","article-title":"The Confusion Effect\u2014From Neural Networks to Reduced Predation Risk","volume":"19","author":"Ioannou","year":"2008","journal-title":"Behav. Ecol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"eabm7548","DOI":"10.1126\/sciadv.abm7548","article-title":"Group Size Affects Predation Risk and Foraging Success in Pacific Salmon at Sea","volume":"8","author":"Polyakov","year":"2022","journal-title":"Sci. Adv."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"708","DOI":"10.1016\/j.cub.2021.11.068","article-title":"Fish Waves as Emergent Collective Antipredator Behavior","volume":"32","author":"Doran","year":"2022","journal-title":"Curr. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1007\/s10164-021-00696-0","article-title":"Dendropsophus Minutus (Hylidae) Tadpole Evaluation of Predation Risk by Fishing Spiders (Thaumasia Sp.: Pisauridae) Is Modulated by Size and Social Environment","volume":"39","author":"Nomura","year":"2021","journal-title":"J. Ethol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1111\/eth.13242","article-title":"Effects of Group Behavior in the Predatory Raid on Damselfish Nests by the False Cleanerfish Aspidontus taeniatus","volume":"128","author":"Sato","year":"2022","journal-title":"Ethology"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1077","DOI":"10.1006\/anbe.1996.0109","article-title":"Why Individual Vigilance Declines as Group Size Increases","volume":"51","author":"Roberts","year":"1996","journal-title":"Anim. Behav."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"104919","DOI":"10.1016\/j.beproc.2023.104919","article-title":"Disentangling the \u201cMany-Eyes\u201d, \u201cDilution Effect\u201d, \u201cSelfish Herd\u201d, and \u201cDistracted Prey\u201d Hypotheses in Shaping Alert and Flight Initiation Distance in a Colonial Seabird","volume":"210","author":"Hammer","year":"2023","journal-title":"Behav. Process."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Zhao, M., Feng, G., Wang, H., Shen, C., Fu, Y., Zhang, Y., Zhang, H., Yao, Y., Chen, J., and Xu, W. (2024). The Influence of Shelter Type and Coverage on Crayfish (Procambarus clarkii) Predation by Catfish (Silurus asotus): A Controlled Environment Study. Animals, 14.","DOI":"10.3390\/ani14081147"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1080\/02705060.2020.1816227","article-title":"Influence of Water Temperature, Habitat Complexity and Light on the Predatory Performance of the Dark Sleeper Odontobutis potamophila (G\u00fcnther, 1861)","volume":"35","author":"Ren","year":"2020","journal-title":"J. Freshwater Ecol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/j.anbehav.2008.01.016","article-title":"Minnows and the Selfish Herd: Effects of Predation Risk on Shoaling Behaviour Are Dependent on Habitat Complexity","volume":"76","author":"Orpwood","year":"2008","journal-title":"Anim. Behav."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"225","DOI":"10.3354\/meps329225","article-title":"Complex Habitats May Not Always Benefit Prey: Linking Visual Field with Reef Fish Behavior and Distribution","volume":"329","author":"Rilov","year":"2007","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Fakan, E.P., Allan, B.J.M., Illing, B., Hoey, A.S., and McCormick, M.I. (2023). Habitat Complexity and Predator Odours Impact on the Stress Response and Antipredation Behaviour in Coral Reef Fish. PLoS ONE, 18.","DOI":"10.1371\/journal.pone.0286570"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e14354","DOI":"10.1111\/rec.14354","article-title":"The Role of Habitat in Predator\u2013Prey Dynamics with Applications to Restoration","volume":"33","author":"Lennox","year":"2025","journal-title":"Restor. Ecol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"240760","DOI":"10.1098\/rsos.240760","article-title":"The Impact of Predators and Vegetation on Shoaling in Wild Zebrafish","volume":"11","author":"Mukherjee","year":"2024","journal-title":"R. Soc. Open Sci."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Encel, S.A., Schaerf, T.M., Lizier, J.T., and Ward, A.J. (2021). Locomotion, Interactions and Information Transfer Vary According to Context in a Cryptic Fish Species. Behav. Ecol. Sociobiol., 75.","DOI":"10.1007\/s00265-020-02930-0"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Liu, L., Zhang, R., Wang, X., Zhu, H., and Tian, Z. (2020). Transcriptome Analysis Reveals Molecular Mechanisms Responsive to Acute Cold Stress in the Tropical Stenothermal Fish Tiger Barb (Puntius tetrazona). BMC Genom., 21.","DOI":"10.1186\/s12864-020-07139-z"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Ramos, T.P.A., Lustosa-Costa, S.Y., Lima, R.M.O., Barbosa, J.E.D.L., and Menezes, R.F. (2021). First Record of Moenkhausia Costae (Steindachner 1907) in the Para\u00edba Do Norte Basin after the S\u00e3o Francisco River Diversion. Biota Neotrop., 21.","DOI":"10.1590\/1676-0611-bn-2020-1049"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1002\/aqc.3761","article-title":"Hatchery Release Programme Modified the Genetic Diversity and Population Structure of Wild Chinese Sucker (Myxocyprinus asiaticus) in the Upper Yangtze River","volume":"32","author":"Wu","year":"2022","journal-title":"Aquat. Conserv. Mar. Freshwater Ecosyst."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"e11434","DOI":"10.1002\/ece3.11434","article-title":"Role of Body Size and Shape in Animal Camouflage","volume":"14","author":"Yu","year":"2024","journal-title":"Ecol. Evol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Degani, G. (2020). Brain Control Reproduction by the Endocrine System of Female Blue Gourami (Trichogaster trichopterus). Biology, 9.","DOI":"10.3390\/biology9050109"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"4387","DOI":"10.1111\/are.14796","article-title":"Physiological Responses to Salinity Stress in the Managua Cichlid, Cichlasoma managuense","volume":"51","author":"Ai","year":"2020","journal-title":"Aquacult. Res."},{"key":"ref_24","unstructured":"(2001). Laboratory Animal\u2014Requirements of Environment and Housing Facilities (Standard No. GB\/T14925-2001)."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1038\/nmeth.2994","article-title":"idTracker: Tracking Individuals in a Group by Automatic Identification of Unmarked Animals","volume":"11","author":"Hinz","year":"2014","journal-title":"Nat. Methods"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1038\/s41592-018-0295-5","article-title":"Idtracker. Ai: Tracking All Individuals in Small or Large Collectives of Unmarked Animals","volume":"16","author":"Bergomi","year":"2019","journal-title":"Nat. Methods"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"2862","DOI":"10.1016\/j.cub.2017.08.004","article-title":"Consistent Individual Differences Drive Collective Behavior and Group Functioning of Schooling Fish","volume":"27","author":"Jolles","year":"2017","journal-title":"Curr. Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"165","DOI":"10.3354\/ab00667","article-title":"Effects of Group Size on Schooling Behavior in Two Cyprinid Fish Species","volume":"25","author":"Fu","year":"2016","journal-title":"Aquat. Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1007\/s11160-012-9260-z","article-title":"Shoals and Schools: Back to the Heuristic Definitions and Quantitative References","volume":"22","author":"Delcourt","year":"2012","journal-title":"Rev. Fish Biol. Fish."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e3222","DOI":"10.7717\/peerj.3222","article-title":"The Behavioral Response of Prey Fish to Predators: The Role of Predator Size","volume":"5","author":"Tang","year":"2017","journal-title":"PeerJ"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Li, W., Li, J., and Fu, S. (2024). Effects of Starve and Shelter Availability on the Group Behavior of Two Freshwater Fish Species (Chindongo demasoni and Spinibarbus sinensis). Animals, 14.","DOI":"10.3390\/ani14162429"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"20220059","DOI":"10.1098\/rstb.2022.0059","article-title":"A Multi-Scale Review of the Dynamics of Collective Behaviour: From Rapid Responses to Ontogeny and Evolution","volume":"378","author":"Ioannou","year":"2023","journal-title":"Phil. Trans. R. Soc. B"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"20190891","DOI":"10.1098\/rspb.2019.0891","article-title":"Inferring Collective Behaviour from a Fossilized Fish Shoal","volume":"286","author":"Mizumoto","year":"2019","journal-title":"Proc. R. Soc. B"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1007\/s10071-018-1229-4","article-title":"Numerical Ability in Fish Species: Preference between Shoals of Different Sizes Varies among Singletons, Conspecific Dyads and Heterospecific Dyads","volume":"22","author":"Bai","year":"2019","journal-title":"Anim. Cogn."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Nadler, L.E., McCormick, M.I., Johansen, J.L., and Domenici, P. (2021). Social Familiarity Improves Fast-Start Escape Performance in Schooling Fish. Commun. Biol., 4.","DOI":"10.1038\/s42003-021-02407-4"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Demandt, N., Bierbach, D., Kurvers, R.H.J.M., Krause, J., Kurtz, J., and Scharsack, J.P. (2021). Parasite Infection Impairs the Shoaling Behaviour of Uninfected Shoal Members under Predator Attack. Behav. Ecol. Sociobiol., 75.","DOI":"10.1007\/s00265-021-03080-7"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"130337","DOI":"10.1016\/j.physa.2024.130337","article-title":"Modeling and Simulation of the Fish Collective Behavior with Risk Perception and Startle Cascades","volume":"659","author":"Zhao","year":"2025","journal-title":"Phys. A"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Horka, P., and Vlachova, M. (2023). The Effect of Turbidity on the Behavior of Bleak (Alburnus alburnus). Fishes, 9.","DOI":"10.3390\/fishes9010003"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e17547","DOI":"10.7717\/peerj.17547","article-title":"Behavioral Responses of a Clonal Fish to Perceived Predation Risk","volume":"12","author":"Jin","year":"2024","journal-title":"PeerJ"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Larsson, M. (2024). Schooling Fish from a New, Multimodal Sensory Perspective. Animals, 14.","DOI":"10.3390\/ani14131984"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"e71798","DOI":"10.1002\/ece3.71798","article-title":"Visual Lateralisation in Female Guppies Poecilia reticulata Demonstrates Social Conformity but Is Reduced When Observing a Live Predator Andinoacara pulcher","volume":"15","author":"Brown","year":"2025","journal-title":"Ecol. Evol."},{"key":"ref_42","first-page":"639","article-title":"Does Social Context Affect Boldness in Juveniles?","volume":"63","author":"Loftus","year":"2017","journal-title":"Curr. Zool."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Li, X., Zhu, Y., Chen, S., Zhu, T., Wu, X., and Li, X. (2025). Behavioral Characteristics of Largefin Longbarbel Catfish Hemibagrus macropterus: Effects of Sex and Body Size on Aggression and Shelter Selection. Animals, 15.","DOI":"10.3390\/ani15091192"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"737930","DOI":"10.1016\/j.aquaculture.2022.737930","article-title":"Effects of Simple Shelters on Growth Performance and Welfare of Rainbow Trout Juveniles","volume":"551","author":"Gesto","year":"2022","journal-title":"Aquaculture"},{"key":"ref_45","first-page":"100908","article-title":"Selection of Shelter Shape by Swimming Crab (Portunus trituberculatus)","volume":"21","author":"Zhang","year":"2021","journal-title":"Aquacult. Rep."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Zhang, H., Zhu, B., Yu, L., and Wang, F. (2022). Shelter Color Selection of Juvenile Swimming Crabs (Portunus trituberculatus). Fishes, 7.","DOI":"10.3390\/fishes7050296"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"802","DOI":"10.1111\/brv.12823","article-title":"Sociability as a Personality Trait in Animals: Methods, Causes and Consequences","volume":"97","author":"Gartland","year":"2022","journal-title":"Biol. Rev."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Zhou, W., Fu, S., Nadler, L.E., Killen, S.S., Zhou, K.-Y., Zheng, S.-L., and Fu, C. (2025). Social Conditions Alter the Growth, Metabolic Rate, Behavioral Traits, and Food-Shelter Preferences in the Juvenile Qingbo Spinibarbus sinensis. Behav. Ecol. Sociobiol., 79.","DOI":"10.1007\/s00265-025-03588-2"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"241","DOI":"10.3354\/meps09264","article-title":"Habitat Type and Schooling Interactively Determine Refuge-Seeking Behavior in a Coral Reef Fish throughout Ontogeny","volume":"437","author":"Huijbers","year":"2011","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1016\/j.applanim.2012.09.005","article-title":"Shelter Use of Familiar and Unfamiliar Groups of Juvenile European Catfish Silurus glanis","volume":"142","author":"Maciak","year":"2012","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1212","DOI":"10.1126\/science.1218919","article-title":"Predatory Fish Select for Coordinated Collective Motion in Virtual Prey","volume":"337","author":"Ioannou","year":"2012","journal-title":"Science"}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/23\/3447\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,4]],"date-time":"2025-12-04T05:19:07Z","timestamp":1764825547000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/23\/3447"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,29]]},"references-count":51,"journal-issue":{"issue":"23","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["ani15233447"],"URL":"https:\/\/doi.org\/10.3390\/ani15233447","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,29]]}}}