{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T10:11:04Z","timestamp":1789035064938,"version":"build-2803163510"},"reference-count":59,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T00:00:00Z","timestamp":1788998400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Insects"],"abstract":"<jats:p>Toxic baits are central to professional ant management, but their efficacy depends on matching macronutrient composition to target species\u2019 preferences, which may vary within species and season. Comparative data across co-occurring Mediterranean ant pests are scarce. We quantified protein, carbohydrate, and mixed-diet preferences in six Mediterranean ant pests\u2014Crematogaster scutellaris, Pheidole pallidula, Tapinoma darioi, Linepithema humile, Lasius neglectus, and Lasius grandis\u2014using 10 replicates per species and season across autumn and spring. Distinct patterns of bait attractiveness were observed among the six studied ant species. Crematogaster scutellaris, P. pallidula, and T. darioi favored protein, whereas invasive L. humile and L. neglectus preferred carbohydrates. Lasius grandis showed no clear preference. Seasonal shifts occurred in some species: P. pallidula and L. neglectus increased protein recruitment in spring, L. grandis favored mixed diets, and L. humile shifted further toward carbohydrates. Thus, carbohydrate-rich matrices are indicated for invasive L. humile and L. neglectus, whereas protein-biased baits suit C. scutellaris, P. pallidula, and T. darioi. Seasonal adjustments, such as increased protein in spring for P. pallidula, may enhance bait attractiveness. These findings provide a species-resolved nutritional framework to guide the development of targeted, season-adapted bait formulations for Mediterranean ant pest management.<\/jats:p>","DOI":"10.3390\/insects17090945","type":"journal-article","created":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T09:13:03Z","timestamp":1789031583000},"page":"945","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Seasonal Variation in Bait Attractiveness Among Six Mediterranean Pest Ant Species: Implications for Toxic Bait Development"],"prefix":"10.3390","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1298-7597","authenticated-orcid":false,"given":"S\u00edlvia","family":"Abril","sequence":"first","affiliation":[{"name":"Departament de Ci\u00e8ncies Ambientals, Universitat de Girona, C. Maria Aur\u00e8lia Capmany 69, 17003 Girona, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c0lex","family":"L\u00f3pez","sequence":"additional","affiliation":[{"name":"Departament de Ci\u00e8ncies Ambientals, Universitat de Girona, C. Maria Aur\u00e8lia Capmany 69, 17003 Girona, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5967-290X","authenticated-orcid":false,"given":"Mara","family":"Moreno-G\u00f3mez","sequence":"additional","affiliation":[{"name":"Research and Development (R&D) Insect Control Department, Henkel Ib\u00e9rica S.A, 08005 Barcelona, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,10]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1093\/jis\/14.1.48","article-title":"Food preference and foraging activity of ants: Recommendations for field applications of low-toxicity baits","volume":"14","author":"Nyamukondiwa","year":"2014","journal-title":"J. Insect Sci."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Wilson, E.O., and H\u00f6lldobler, B. (1990). The Ants, Harvard University Press.","DOI":"10.1007\/978-3-662-10306-7"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Bl\u00fcthgen, N., and Feldhaar, H. (2010). Food and shelter: How resources influence ant ecology. Ant Ecology, Oxford University Press.","DOI":"10.1093\/acprof:oso\/9780199544639.003.0007"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1096","DOI":"10.1111\/1744-7917.13461","article-title":"Foraging by predatory ants: A review","volume":"32","author":"Dejean","year":"2025","journal-title":"Insect Sci."},{"key":"ref_5","first-page":"317","article-title":"Laboratory evaluation of baits, residual insecticides, and an ultrasonic device for control of white-footed ants Technomyrmex albipes (Hymenoptera: Formicidae)","volume":"45","author":"Warner","year":"2005","journal-title":"Sociobiology"},{"key":"ref_6","unstructured":"Corsetti, K.A. (2024). Bait Acceptance and Seasonal Activity of Brachyponera chinensis. [Master\u2019s Thesis, University of Georgia]."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"5727","DOI":"10.1038\/s41598-024-56133-y","article-title":"Effects of macro- and micro-nutrients on momentary and season-long feeding responses by select species of ants","volume":"14","author":"Renyard","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1098\/rstb.1993.0166","article-title":"A multi-level analysis of feeding behaviour: The geometry of nutritional decisions","volume":"342","author":"Simpson","year":"1993","journal-title":"Phil. Trans. R. Soc. B"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1016\/j.anbehav.2009.11.022","article-title":"Colony-level macronutrient regulation in ants: Mechanisms, hoarding and associated costs","volume":"79","author":"Cook","year":"2010","journal-title":"Anim. Behav."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1016\/j.anbehav.2015.10.021","article-title":"Responses to nutritional challenges in ant colonies","volume":"111","author":"Bazazi","year":"2016","journal-title":"Anim. Behav."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1007\/s00114-010-0705-8","article-title":"Nutrient compensatory foraging in a free-living social insect","volume":"97","author":"Christensen","year":"2010","journal-title":"Naturwissenschaften"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1111\/j.1744-7429.2009.00616.x","article-title":"Macronutrient regulation in the tropical terrestrial ant Ectatomma ruidum (Formicidae): A field study in Costa Rica","volume":"42","author":"Cook","year":"2010","journal-title":"Biotropica"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1238","DOI":"10.1093\/aesa\/63.5.1238","article-title":"The Seasonal life cycle of the Argentine ant, Iridomyrmex humilis (Hymenoptera: Formicidae), in Southern California","volume":"63","author":"Markin","year":"1970","journal-title":"Ann. Entomol. Soc. Am."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1093\/jee\/63.3.740","article-title":"Foraging behavior of the Argentine ant in a California citrus grove","volume":"63","author":"Markin","year":"1970","journal-title":"J. Econ. Entomol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1603\/0046-225X(2007)36[1166:FAADSO]2.0.CO;2","article-title":"Foraging activity and dietary spectrum of the Argentine ant (Hymenoptera: Formicidae) in invaded natural areas of the northeast Iberian Peninsula","volume":"36","author":"Abril","year":"2007","journal-title":"Environ. Entomol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1016\/j.cub.2009.03.015","article-title":"Communal nutrition in ants","volume":"19","author":"Dussutour","year":"2009","journal-title":"Curr. Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"117","DOI":"10.2307\/3495334","article-title":"Seasonal changes in bait preference by red imported fire ant, Solenopsis invicta (Hymenoptera: Formicidae)","volume":"73","author":"Stein","year":"1990","journal-title":"Fla. Entomol."},{"key":"ref_18","unstructured":"Barbani, L.E. (2003). Foraging activity and food preferences of the odorous house ant Tapinoma sessile. [Master\u2019s Thesis, Virginia Polytechnic Institute and State University]."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.jinsphys.2016.02.001","article-title":"Summer and fall ants have different physiological responses to food macronutrient content","volume":"87","author":"Cook","year":"2016","journal-title":"J. Insect Physiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1093\/aesa\/90.2.246","article-title":"Food preference and patterns of foraging activity of the southern fire ant (Hymenoptera: Formicidae)","volume":"90","author":"Hooper","year":"1997","journal-title":"Ann. Entomol. Soc. Am."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1093\/jee\/100.2.509","article-title":"Relative attractiveness of baits to Paratrechina longicornis (Hymenoptera: Formicidae)","volume":"100","author":"Stanley","year":"2007","journal-title":"J. Econ. Entomol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1482","DOI":"10.1093\/ee\/27.6.1482","article-title":"Bait preference by the Argentine ant (Hymenoptera: Formicidae) in Haleakal\u0101 National Park, Hawaii","volume":"27","author":"Krushelnycky","year":"1998","journal-title":"Environ. Entomol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1603\/0046-225X-31.6.977","article-title":"Patterns of macronutrient collection in the black carpenter ant Camponotus pennsylvanicus (De Geer) (Hymenoptera: Formicidae)","volume":"31","author":"Cannon","year":"2002","journal-title":"Environ. Entomol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"802","DOI":"10.1093\/ee\/16.3.802","article-title":"Foraging in Solenopsis Invicta (Hymenoptera: Formicidae): Effects of weather and season","volume":"16","author":"Porter","year":"1987","journal-title":"Environ. Entomol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"19","DOI":"10.2980\/20-1-3559","article-title":"A year in an ant\u2019s life: Opportunism and seasonal variation in the foraging ecology of Aphaenogaster senilis","volume":"20","author":"Caut","year":"2013","journal-title":"Ecoscience"},{"key":"ref_26","first-page":"123","article-title":"Four species within the supercolonial ants of the Tapinoma nigerrimum complex revealed by integrative taxonomy (Hymenoptera: Formicidae)","volume":"24","author":"Seifert","year":"2017","journal-title":"Myrmecol. News"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"434","DOI":"10.1111\/jen.12751","article-title":"Six new non-native ants (Formicidae) in the Canary Islands and their possible impacts","volume":"144","year":"2020","journal-title":"J. Appl. Entomol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1890\/0012-9658(1999)080[0238:CMUTDO]2.0.CO;2","article-title":"Competitive mechanisms underlying displacement of native ants by the invasive Argentine ant","volume":"80","author":"Holway","year":"1999","journal-title":"Ecology"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1095","DOI":"10.1073\/pnas.98.3.1095","article-title":"Patterns of spread in biological invasions dominated by long-distance jump dispersal: Insights from Argentine ants","volume":"98","author":"Suarez","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1127\/entomologia\/2023\/2187","article-title":"The Argentine ant Linepithema humile: Natural history, ecology and impact of a successful invader","volume":"44","author":"Angulo","year":"2024","journal-title":"Entomol. Gen."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Cremer, S., Ugelvig, L.V., Drijfhout, F.P., Schlick-Steiner, B.C., Steiner, F.M., Seifert, B., Hughes, D.P., Schulz, A., Petersen, K.S., and Konrad, H. (2008). The evolution of invasiveness in garden ants. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0003838"},{"key":"ref_32","first-page":"99","article-title":"Area-wide management of the invasive garden ant Lasius neglectus (Hymenoptera: Formicidae) in Northeast Spain","volume":"21","author":"Rey","year":"2004","journal-title":"J. Agric. Urban Entomol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1111\/een.13395","article-title":"New crop, new pest, old ants: Crop colonisation by native ants disrupt biological control of an invasive mealybug in Mediterranean persimmon","volume":"50","author":"Plata","year":"2025","journal-title":"Ecol. Entomol."},{"key":"ref_34","first-page":"249","article-title":"Ant species noxious to cork oak in Sardinia","volume":"76","author":"Verdinelli","year":"2024","journal-title":"IOBC\/WPRS Bull."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/s13595-015-0534-1","article-title":"Cork oak pests: A Review of insect damage and management: R. Tiberi et al.","volume":"73","author":"Tiberi","year":"2016","journal-title":"Ann. For. Sci."},{"key":"ref_36","unstructured":"(2026, March 06). Global Invasive Species Database (GISD). Available online: https:\/\/www.iucngisd.org\/gisd\/100_worst.php."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"6075","DOI":"10.1073\/pnas.092694199","article-title":"Evolution of supercolonies of Argentine ants of Southern Europe","volume":"99","author":"Giraud","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_38","unstructured":"Arcos, J., and Garc\u00eda-Garc\u00eda, F. (2023). Hormigas de la Pen\u00ednsula Ib\u00e9rica e Islas Baleares, Independently Published."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1603\/022.038.0203","article-title":"Ant community structure in citrus orchards in the Mediterranean Basin: Impoverishment as a consequence of habitat homogeneity","volume":"38","author":"Palacios","year":"2009","journal-title":"Environ. Entomol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1155\/1991\/75196","article-title":"Dynamics of collective exploration in the ant Pheidole pallidula","volume":"98","author":"Detrain","year":"1991","journal-title":"Psyche"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"229","DOI":"10.2307\/3493193","article-title":"Artificial diet for rearing ants","volume":"53","author":"Bhatkar","year":"1970","journal-title":"Fla. Entomol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1007\/s00040-008-1008-3","article-title":"Description of a simple synthetic diet for studying nutritional responses in ants","volume":"55","author":"Dussutour","year":"2008","journal-title":"Insectes Sociaux"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.scitotenv.2019.05.260","article-title":"Food resource exploitation and functional resilience in ant communities found in common Mediterranean habitats","volume":"684","author":"Arnan","year":"2019","journal-title":"Sci. Total Environ."},{"key":"ref_44","unstructured":"R Core Team (2018). R: A Language and Environment for Statistical Computing, R Foundation for Statistical Computing."},{"key":"ref_45","first-page":"637","article-title":"Estudio del comportamiento alimentario de Crematogaster scutellaris Oliv. (Hym. Formicidae) en tres alcornocales del SW espa\u00f1ol","volume":"20","year":"1994","journal-title":"Bol. San. Veg. Plagas"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1007\/s11829-009-9057-8","article-title":"Honeydew collection by the invasive garden ant Lasius neglectus versus the native ant L. grandis","volume":"3","author":"Paris","year":"2009","journal-title":"Arthropod-Plant Interact."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1016\/j.cropro.2011.03.014","article-title":"Preference of foraging ants (Hymenoptera: Formicidae) for bait toxicants in South African vineyards","volume":"30","author":"Nyamukondiwa","year":"2011","journal-title":"Crop Prot."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1007\/BF01907668","article-title":"Acceptability of different sugars and oils to three tropical ant species (Hymenoptera: Formicidae)","volume":"69","author":"Cornelius","year":"1996","journal-title":"Anz. Sch\u00e4dlingskde. Pflanzenschutz Umweltschutz"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2627","DOI":"10.1002\/ps.5365","article-title":"Previous diet affects the amount but not the type of bait consumed by an invasive ant","volume":"75","author":"Lach","year":"2019","journal-title":"Pest Manag. Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"684","DOI":"10.1111\/een.12146","article-title":"Carbohydrate scarcity increases foraging activities and aggressiveness in the ant Prolasius advenus (Hymenoptera: Formicidae)","volume":"39","author":"Grangier","year":"2014","journal-title":"Ecol. Entomol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2951","DOI":"10.1098\/rspb.2007.1065","article-title":"Linking nutrition and behavioural dominance: Carbohydrate scarcity limits aggression and activity in Argentine ants","volume":"274","author":"Grover","year":"2007","journal-title":"Proc. R. Soc. B"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1890\/ES11-00145.1","article-title":"Trait-mediated indirect interactions in invasions: Unique behavioral responses of an invasive ant to plant nectar","volume":"2","author":"Savage","year":"2011","journal-title":"Ecosphere"},{"key":"ref_53","first-page":"29","article-title":"The enigma of sugar: Changes in aggressiveness of weaver ants across habitats in response to sugar supplies","volume":"36","author":"Zhang","year":"2026","journal-title":"Myrmecol. News"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Horn, K.C., Eubanks, M.D., and Siemann, E. (2013). The effect of diet and opponent size on aggressive interactions involving Caribbean crazy ants (Nylanderia fulva). PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0066912"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1111\/j.1365-3032.1989.tb00947.x","article-title":"Queen execution in Argentine ants, Iridomyrmex humilis","volume":"14","author":"Keller","year":"1989","journal-title":"Physiol. Entomol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.jinsphys.2014.01.004","article-title":"Strength in numbers: Large and permanent colonies have higher queen oviposition rates in the invasive Argentine ant (Linepithema humile, Mayr)","volume":"62","author":"Abril","year":"2014","journal-title":"J. Insect Physiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1007\/BF02223196","article-title":"Incidence des facteurs \u00e9cologiques sur le cycle annuel et l\u2019activit\u00e9 saisonni\u00e8re de la fourmi d\u2019Argentine, Iridomyrmex humilis Mayr (Hymenoptera, Formicidae), dans la r\u00e9gion d\u2019Antibes","volume":"20","author":"Benois","year":"1973","journal-title":"Insectes Sociaux"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Abril, S., D\u00edaz, M., Lenoir, A., Paris, C.I., Boulay, R., and G\u00f3mez, C. (2018). Cuticular hydrocarbons correlate with queen reproductive status in native and invasive Argentine ants (Linepithema humile, Mayr). PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0193115"},{"key":"ref_59","first-page":"30","article-title":"Food and bait preferences of Liometopum occidentale (Hymenoptera: Formicidae)","volume":"49","author":"Rust","year":"2014","journal-title":"J. Entomol. Sci."}],"container-title":["Insects"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-4450\/17\/9\/945\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T09:36:14Z","timestamp":1789032974000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-4450\/17\/9\/945"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,10]]},"references-count":59,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["insects17090945"],"URL":"https:\/\/doi.org\/10.3390\/insects17090945","relation":{},"ISSN":["2075-4450"],"issn-type":[{"value":"2075-4450","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,10]]}}}