{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,2]],"date-time":"2026-01-02T07:46:41Z","timestamp":1767340001584,"version":"build-2065373602"},"reference-count":43,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2023,1,12]],"date-time":"2023-01-12T00:00:00Z","timestamp":1673481600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"China Scholarship Council","award":["2017-70006-27290"],"award-info":[{"award-number":["2017-70006-27290"]}]},{"name":"USDA","award":["2017-70006-27290"],"award-info":[{"award-number":["2017-70006-27290"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Insects"],"abstract":"<jats:p>Numerous studies have confirmed that the trade-off between anti-predator behavior and mating behavior occurs in certain insect species. This suggests that insects invest more in anti-predator behavior, and fewer resources or time can be used in mating behavior. However, few studies focus on tonic immobility, an important anti-predator behavior in nature, and different stages in mating behavior. Tonic immobility (TI) is considered to be an important anti-predator behavior. Herein, we investigated the relationship between TI and mating behavior in the sweetpotato weevil (SPW), Cylas formicarius. As the first step, we artificially selected SPWs for the longer duration of TI (L-strain) and the shorter duration of TI (S-strain). The effect of courtship and copulation on the duration of TI in two artificial selection strains was tested. Furthermore, we compared the frequency and duration of two mating behaviors in four kinds of pairs (LF\u00d7LM, LF\u00d7SM, SF\u00d7LM, and SF\u00d7SM: LM\u2014L-strain male; SM\u2014S-strain male; LF\u2014L-strain female; SF\u2014S-strain female). Finally, we tested insemination success in four kinds of pairs (male and female SPWs from the L-strain or the S-strain). The courtship and copulation significantly reduced the duration of TI. Pairs with males from the L-strain showed lower frequency and longer duration of courtship than pairs with males from the S-strain. Similarly, males from L-strain pairs showed a longer period of copulation than pairs with males from the S-strain. However, there is no significant difference in the frequency of copulation and the success of insemination. These results support that there was a significant trade-off between TI and courtship as well as copulation in the SPW.<\/jats:p>","DOI":"10.3390\/insects14010073","type":"journal-article","created":{"date-parts":[[2023,1,12]],"date-time":"2023-01-12T03:47:01Z","timestamp":1673495221000},"page":"73","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Investment Trade-Off between Mating Behavior and Tonic Immobility in the Sweetpotato Weevil Cylas formicarius (Coleoptera: Brentidae)"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7927-1094","authenticated-orcid":false,"given":"Haoyong","family":"Ouyang","sequence":"first","affiliation":[{"name":"State Key Laboratory of Integrated Management of Pest Insects and Rodents, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China"},{"name":"College of Life Science, University of Chinese Academy of Sciences, No. 19(A) Yuquan Road, Shijingshan District, Beijing 100049, China"},{"name":"Center for Biological Control, College of Agriculture and Food Sciences, Florida A&M University, Tallahassee, FL 32307, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9001-0154","authenticated-orcid":false,"given":"Runzhi","family":"Zhang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Integrated Management of Pest Insects and Rodents, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China"},{"name":"College of Life Science, University of Chinese Academy of Sciences, No. 19(A) Yuquan Road, Shijingshan District, Beijing 100049, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2208-5984","authenticated-orcid":false,"given":"Muhammad","family":"Haseeb","sequence":"additional","affiliation":[{"name":"Center for Biological Control, College of Agriculture and Food Sciences, Florida A&M University, Tallahassee, FL 32307, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2023,1,12]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1007\/BF00345569","article-title":"Coexistence of two species of whip spiders (Genus Heterophrynus) in the Neotropical rain forest (Arachnida, Amblypygi)","volume":"27","author":"Weygoldt","year":"1977","journal-title":"Oecologia"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"80","DOI":"10.5962\/p.266539","article-title":"Phylogenetic, habitat, and behavioural aspects of possum behaviour in European Lepidoptera","volume":"39","author":"Shreeve","year":"2006","journal-title":"J. Res. Lepid."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"139","DOI":"10.2307\/3492753","article-title":"Some notes on the citrus root weevil","volume":"35","author":"Wolfenbarger","year":"1952","journal-title":"Fla. Entomol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1007\/BF00295144","article-title":"Mammalian Breath: Trigger of defensive chemical response in a Tenebrionid beetle (Bolitotherus Cornutus)","volume":"16","author":"Conner","year":"1985","journal-title":"Behav. Ecol. Sociobiol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1023\/A:1011196420147","article-title":"Diurnal Periodicity of death-feigning in Cylas formicarius (Coleoptera: Brentidae)","volume":"14","author":"Miyatake","year":"2001","journal-title":"J. Insect Behav."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"429","DOI":"10.2307\/3494197","article-title":"The larva of Neurocordulia Xanthosoma (Odonata: Corduliidae)","volume":"59","author":"Williams","year":"1976","journal-title":"Fla. Entomol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1007\/s00114-008-0362-3","article-title":"Young fire ant workers feign death and survive aggressive neighbors","volume":"95","author":"Cassill","year":"2008","journal-title":"Naturwissenschaften"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1016\/0003-3472(60)90025-7","article-title":"Immobility reactions (fear) of domestic fowl as a function of age and prior experience","volume":"8","author":"Ratner","year":"1960","journal-title":"Anim. Behav."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"108","DOI":"10.2307\/2424542","article-title":"Death feigning by ducks in response to predation by red foxes (Vulpes Fulva)","volume":"94","author":"Sargeant","year":"1975","journal-title":"Am. Midl. Nat."},{"key":"ref_10","first-page":"135","article-title":"Thanatosis in four poorly known toads of the Genus Incilius (Amphibia: Anura) from the highlands of costa rica","volume":"3","author":"Abarca","year":"2016","journal-title":"Mesoam Herpetol"},{"key":"ref_11","first-page":"78","article-title":"death-feigning behavior of the Japanese colubrid snake Rhabdophis tigrinus","volume":"39","author":"Mutoh","year":"1983","journal-title":"Herpetologica"},{"key":"ref_12","first-page":"89","article-title":"Predator simulation and duration of death feigning in neonate hognose snakes","volume":"7","author":"Burghardt","year":"1977","journal-title":"Anim. Behav."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1146\/annurev.ecolsys.31.1.79","article-title":"The evolution of predator-prey interactions: Theory and evidence","volume":"31","author":"Abrams","year":"2000","journal-title":"Annu. Rev. Ecol. Syst"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Sakai, M. (2021). Death-Feigning in Insects Mechanism and Function of Tonic Immobility, Springer.","DOI":"10.1007\/978-981-33-6598-8"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"847","DOI":"10.1086\/675903","article-title":"When to defend: Antipredator defenses and the predation sequence","volume":"183","author":"Bateman","year":"2014","journal-title":"Am. Nat."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Choe, J.C., and Crespi, B.J. (1997). Evolutionary perspectives on insect mating. The Evolution of Mating Systems in Insects and Arachnids, Cambridge University Press.","DOI":"10.1017\/CBO9780511721946"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1093\/beheco\/ary144","article-title":"Male courtship behaviors and female choice reduced during experimental starvation stress","volume":"30","author":"Billings","year":"2019","journal-title":"Behav. Ecol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/j.exger.2010.10.008","article-title":"Survival Costs of reproduction in Drosophila","volume":"46","author":"Flatt","year":"2011","journal-title":"Exp. Gerontol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1016\/S0022-1910(03)00117-3","article-title":"Longevity cost of reproduction for males but no longevity cost of mating or courtship for females in the male-dimorphic dung beetle Onthophagus Binodis","volume":"49","author":"Kotiaho","year":"2003","journal-title":"J. Insect Physiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"955","DOI":"10.2307\/2410417","article-title":"Resource allocation and mating systems in butterflies","volume":"49","author":"Karlsson","year":"1995","journal-title":"Evolution"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1262","DOI":"10.1111\/j.1365-2435.2010.01756.x","article-title":"Experimental evidence for physiological costs underlying the trade-off between reproduction and survival","volume":"24","author":"Cox","year":"2010","journal-title":"Funct. Ecol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1016\/j.bbagen.2009.07.010","article-title":"Integrating evolutionary and molecular genetics of aging","volume":"1790","author":"Flatt","year":"2009","journal-title":"Biochim. Biophys. Acta Gen. Subj."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1093\/beheco\/arr173","article-title":"Costly Copulation in the Wild: Mating Increases the risk of parasitoid-mediated death in swarming locusts","volume":"23","author":"Kemp","year":"2012","journal-title":"Behav. Ecol."},{"key":"ref_24","first-page":"1","article-title":"The sweetpotato weevils in Hawaii their biology and control","volume":"23","author":"Sherman","year":"1954","journal-title":"Hawaii Agric. Exp. Stn."},{"key":"ref_25","first-page":"1","article-title":"The biology of the sweet potato weevil","volume":"483","author":"Cockerham","year":"1954","journal-title":"La. Agric. Exp. Stn. Tech. Bull."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1016\/j.anbehav.2009.07.031","article-title":"Copulation reduces the duration of death-feigning behaviour in the sweetpotato weevil, Cylas formicarius","volume":"78","author":"Kuriwada","year":"2009","journal-title":"Anim. Behav."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Ouyang, H., Wu, P., Zhang, R., and Haseeb, M. (2020). Trade-off investment between tonic immobility and mate search in the sweetpotato weevil, Cylas formicarius (Coleoptera: Brentidae). Insects, 11.","DOI":"10.3390\/insects11110774"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2293","DOI":"10.1098\/rspb.2004.2858","article-title":"Is Death-Feigning Adaptive? Heritable Variation in fitness difference of death-feigning behaviour","volume":"271","author":"Miyatake","year":"2004","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1098\/rsbl.2009.0494","article-title":"Genetic Trade-off between Abilities to Avoid Attack and to Mate: A Cost of Tonic Immobility","volume":"6","author":"Nakayama","year":"2010","journal-title":"Biol. Lett."},{"key":"ref_30","unstructured":"R Core Team (2022). R: A Language and Environment for Statistical Computing, R Foundation for Statistical Computing."},{"key":"ref_31","unstructured":"Wickham, H., Chang, W., Henry, L., Pedersen, T.L., Takahashi, K., Wilike, C., Woo, K., Yutani, H., and Dunnington, D. (2022, November 23). ggplot2: Create Elegant Data Visualisations Using the Grammar of Graphics. Available online: https:\/\/ggplot2.tidyverse.org\/."},{"key":"ref_32","unstructured":"Bates, D., Maechler, M., Bolker, B., Walker, S., Christensen, R.H.B., Singmann, H., Dai, B., Scheipl, F., Grothendieck, G., and Green, P. (2022, November 23). lme4: Linear Mixed-Effects Models Using \u201cEigen\u201d and S4. Available online: http:\/\/cran.r-project.org\/web\/packages\/lme4\/index.html."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/j.tree.2008.10.008","article-title":"Generalized linear mixed models: A practical guide for ecology and evolution","volume":"24","author":"Bolker","year":"2009","journal-title":"Trends. Ecol. Evol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1054","DOI":"10.1093\/beheco\/arq104","article-title":"Mobbing as a trade-off between safety and reproduction in a songbird","volume":"21","author":"Krama","year":"2010","journal-title":"Behav. Ecol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"848","DOI":"10.1016\/j.jinsphys.2008.03.003","article-title":"Influence of age and juvenile hormone on brain dopamine level in male honeybee (Apis mellifera): Association with reproductive maturation","volume":"54","author":"Harano","year":"2008","journal-title":"J. Insect Physiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"5539","DOI":"10.1523\/JNEUROSCI.5290-07.2008","article-title":"Increased dopamine level enhances male-male courtship in Drosophila","volume":"28","author":"Liu","year":"2008","journal-title":"J. Neurosci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1023\/A:1015279221600","article-title":"Effects of Tyrosine Hydroxylase Mutants on Locomotor Activity in Drosophila: A Study in Functional Genomics","volume":"32","author":"Pendleton","year":"2002","journal-title":"Behav. Genet."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.anbehav.2007.04.019","article-title":"Pleiotropic antipredator strategies, fleeing and feigning death, correlated with dopamine levels in Tribolium castaneum","volume":"75","author":"Miyatake","year":"2008","journal-title":"Anim. Behav."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1086\/589894","article-title":"Sexual Selection for Male Mobility in a Giant Insect with Female-Biased Size Dimorphism","volume":"172","author":"Kelly","year":"2008","journal-title":"Am. Nat."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1163\/156853999501621","article-title":"Sperm Competition in a Nuptial Feeding Spider, Pisaura Mirabilis","volume":"136","author":"Drengsgaard","year":"1999","journal-title":"Behaviour"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1111\/j.1469-185X.2008.00037.x","article-title":"An Integrative View of Sexual Selection in Tribolium Flour Beetles","volume":"83","author":"Fedina","year":"2008","journal-title":"Biol. Rev."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1111\/j.1365-3032.2010.00777.x","article-title":"Age-dependent investment in death-feigning behaviour in the sweetpotato weevil Cylas formicarius","volume":"36","author":"Kuriwada","year":"2011","journal-title":"Physiol. Entomol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1093\/beheco\/arr170","article-title":"Resource allocation trade-off between sperm quality and immunity in the field cricket, Teleogryllus oceanicus","volume":"23","author":"Simmons","year":"2012","journal-title":"Behav. Ecol."}],"container-title":["Insects"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-4450\/14\/1\/73\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T18:03:45Z","timestamp":1760119425000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-4450\/14\/1\/73"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,1,12]]},"references-count":43,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2023,1]]}},"alternative-id":["insects14010073"],"URL":"https:\/\/doi.org\/10.3390\/insects14010073","relation":{},"ISSN":["2075-4450"],"issn-type":[{"type":"electronic","value":"2075-4450"}],"subject":[],"published":{"date-parts":[[2023,1,12]]}}}