Do honey and pollen enhance the reproductive performance of ladybugs? A case study on Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae)

Do honey and pollen enhance the reproductive performance of ladybugs? 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A case study on Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae) Maurício Silva Lima, Wendel José Teles Pontes, Elio Cesar Guzzo, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7160590/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 28 Oct, 2025 Read the published version in International Journal of Tropical Insect Science → Version 1 posted 15 You are reading this latest preprint version Abstract Coccinellids (Coleoptera: Coccinellidae) are omnivorous insects. In addition to their primary prey, which is the essential food, they also consume alternative foods such as pollen, honey, and fungi. Recent studies have suggested that the combination of essential and alternative foods optimizes the performance of coccinellids, with the alternative food acting as a supplement, in the absence of the essential one. The alternative food would be actively sought, even if the essential one was abundant, as the simultaneous consumption of both would improve the predator's performance. Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae) is an important predator, primarily of mealybugs. To test the hypothesis, we carried out an experiment that consisted of three treatments: adult females of T. notata fed only with the mealybug Ferrisia dasylirii (Cockerell, 1896) (Hemiptera: Pseudococcidae); with F. dasylirii + pollen; or with F. dasylirii + bee honey. There was no statistical difference among the three treatments regarding periods of pre-oviposition (5.4, 5.2, and 5.0 days, respectively), oviposition (15.7, 8.8, and 15.7 days, respectively), and post-oviposition (16.4, 21.9, and 16.6 days, respectively), longevity (36.2, 38.0, and 32.3 days, respectively), and fertility (75.3%, 71.4%, and 73.7%, respectively) of T. notata females. Supplementation of the essential food with pollen significantly reduced fecundity (33.7 eggs/female) in relation to honey and the essential food alone (50.9 and 51.8 eggs/female, respectively), which, in turn, did not differ from each other. The results refuted the hypothesis that alternative food would optimize the reproductive performance of T. notata females. Nutricional ecology Insect nutrition Insect feeding Reproductive fitness Full Text Ladybugs, also known as ladybird beetles (Coleoptera: Coccinellidae), are among the most important predators of pests in agricultural crops (Obrycki and Kring 1998; Iperti 1999; Souza et al. 2019; Sujii et al. 2020; Bueno and van Lenteren 2021). These beetles are the primary predators of scale insects (Hemiptera: Diaspididae, Dactylopiidae, and Pseudococcidae) (Lima et al. 2011, 2016, 2018), with which they are often found associated, and are considered their essential food source. The ladybug Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae), in both larval and adult stages, preys on scale insects and is used, for example, as a complement to biological control carried out by the parasitoid Anagyrus lopezi (De Santis, 1964) (Hymenoptera: Encyrtidae) on the mealybug Phenacoccus manihoti Matile-Ferrero, 1977 (Hemiptera: Pseudococcidae) in cassava Manihot esculenta Crantz crops in Africa (Stäubli Dreyer et al. 1997). The efficient mass production of natural enemies is a fundamental tool within Integrated Pest Management, aiming to provide parasitoids and predators with biological and behavioral characteristics compatible with natural populations and capable of reducing the population density of economically important pests while maintaining economic viability in the production system (Zanuncio et al. 2002; Souza and Souza-Bezerra 2019; Bertin et al. 2021; Parra 2021). To achieve this, predatory insects rely on factors such as the quality and quantity of food to mitigate adverse effects on their biological aspects, such as development, reproduction, and longevity (Parajulee and Phillips 1993). Alternative foods, such as pollen, nectar, honeydew, honey, fruits, foliage, and fungi (Pemberton and Vandenberg 1993; Cividanes et al. 2007; Lundgren 2009), can help to maintain predator populations in their habitat in response to changes in pest population density (Lundgren 2009). Although pollen does not play a vital role in the ladybugs' diet, 39 species of ladybugs that consume pollen from 88 plant species have been identified, under both field and laboratory conditions (Lundgren 2009). Pollen can promote spermatogenesis (Hemptinne and Naisse 1987) and, when combined with prey, can aid in the reproduction of certain species (De Clercq et al. 2005; Berkvens et al. 2008). Carbohydrate-rich foods, such as honey, which is rich in glucose and fructose, can significantly improve the adult survival of ladybirds and, when combined with essential food, can allow females to enhance their reproductive performance (Lundgren 2009). Essential foods support the development and reproduction of ladybugs, while alternative foods are helpful as an energy source, prolonging their survival (Hodek and Honek 1996; Almeida and Ribeiro-Costa 2009, 2012). Many studies have proposed that essential food, combined with alternative food, can enhance the performance of ladybugs, serving as a supplement in prey scarcity, as they actively seek these alternative foods even when prey is abundant. Therefore, the simultaneous consumption of both types of food would increase the performance of these predators (Lundgren 2009). However, other studies suggest that dietary supplementation with pollen may have a negative influence on ladybug reproduction (Lima et al. 2020). Thus, the present study aimed to evaluate the effect of dietary supplementation on the biological performance of T. notata females, testing the hypothesis that supplementary food should optimize the biological parameters of this species when compared to individuals exclusively fed on their essential food, the mealybug Ferrisia dasylirii (Cockerell, 1896) (Hemiptera: Pseudococcidae), under laboratory conditions. The experiment was conducted at the Laboratory of Plant Biology and Insect Resistance, located in the Department of Agronomy at the Federal Rural University of Pernambuco (UFRPE), in Recife, Pernambuco, Brazil. Individuals of F. dasylirii and T. notata used in the experiments were obtained from the insect rearing facility provided by the Laboratory of Biological Control and Insect Ecology at UFRPE. The rearing facilities were established in a climate-controlled room with controlled conditions (25 ± 2 °C, 70 ± 10% relative humidity, and a 12-hour photoperiod). To rear F. dasylirii , pumpkins Cucurbita maxima Duch (Cucurbitaceae) cv. Jacarezinho, in its initial stage of ripening, was used as a host plant. All fruits were sanitized with a 10% sodium hypochlorite solution before infestation. The pumpkins were placed on a glass support inside plastic trays (30 × 45 cm), which were in turn placed on a larger plastic tray (64 × 63 cm) containing water + detergent to prevent infestation by ants and other undesirable insects. Infested fruits were placed alongside non-infested fruits, allowing nymphs to move from one to the other. In the absence of nymphs, females were transferred to non-infested fruits using a fine-bristle brush. Plastic cages (40 × 25 × 20 cm) covered with voile were used for rearing T. notata . Infested pumpkins with F. dasylirii were provided as food. To prevent damage to the fruit and its rapid deterioration, the pumpkins were placed on a metal support protected by a sponge. As a supplementary food source, honeybee pollen, 5% honey, or eggs of Ephestia kuehniella Zeller, 1879 (Lepidoptera: Pyralidae) were offered. Young and adult forms were reared separately. Pumpkins were periodically replaced with newly infested fruits or when they became deteriorated. To obtain T. notata individuals of known and standardized age, pairs of the insect from the rearing facility were separated into plastic containers (300 mL) and fed with F. dasylirii adults. Egg masses were removed after 24 hours, and the larvae from these eggs were fed with F. dasylirii nymphs until the emergence of adults, as preliminary tests showed that the larvae are highly specific and do not feed on pollen and honey. Pairs of T. notata adults of the same age were removed from the rearing cage and individually placed in plastic containers (300 mL) containing one of the following treatments. The containers were labeled and kept in a B.O.D. (Biological Oxygen Demand) incubator at a temperature of 25 ± 1 °C, relative humidity of 70 ± 10%, and a 12-hour photophase. Male T. notata were removed from the containers after 72 hours. The experiment consisted of three treatments: 1) females fed only with F. dasylirii ; 2) females fed with F. dasylirii + honey; and 3) females fed with F. dasylirii + pollen. Pure honey (diluted to 5% for the experiments) and commercial honeybee pollen were used. Mealybugs were offered daily ad libitum directly in the container, and honey and pollen were offered, respectively, on moistened cotton and in small plastic containers, and replaced every 3 days until the death of the females. The investigated biological parameters included the pre-oviposition, oviposition, and post-oviposition periods, as well as longevity, fertility, and fecundity. The experimental design was completely randomized, with 20 replications, where each container with a T. notata female was considered as one replication. Means were subjected to analysis of variance and compared using the t-test at a 5% significance level, using the PROC ANOVA function of SAS statistical software (SAS Institute 2001). The results showed that feeding adult female T. notata with nymphs of F. dasylirii , nymphs of F. dasylirii + pollen, or nymphs of F. dasylirii + honey did not cause significant differences in the pre-oviposition (5.4, 5.2, and 5.0 days, respectively) ( F = 0.11; p = 0.89), oviposition (15.7, 8.8, and 15.7 days, respectively) ( F = 2.43; p = 0.09), or post-oviposition (16.4, 21.9, and 16.6 days, respectively) ( F = 0.40; p = 0.67) periods, which was reflected in their longevity (36.2, 38.0, and 32.3 days, respectively) ( F = 0.18; p = 0.83) (Table 1), which encompasses the pre-oviposition, oviposition, and post-oviposition periods combined. Table 1 here Correia and Berti Filho (1988) highlight that complementary feeding serves as an energy source that can only prolong the survival of adult ladybugs. Additionally, Smith and Krischik (1999) suggest that sugar-based feeding can reduce the pre-oviposition period of Coccinellidae and help females to survive during diapause periods. In Africa, pure honey serves as a food source for adult T. notata , which is employed in the biological control of P. manihoti in cassava, allowing ladybugs to maintain their longevity during transportation (Herren and Neuenschwander 1991; Neuenschwander and Zweigert 1994). However, such effects were not observed in the present study and appear not to apply to T. notata and the provided foods. As Lundgren (2009) suggests, alternative food serves to stabilize the predator population during periods of prey scarcity when it diversifies its food type in response to changes in the population density of its essential food source within its niche. However, in the current study, there was no restriction on the natural prey ( F. dasylirii ) for the predator T. notata . Regarding fecundity, T. notata females laid fewer eggs when fed with F. dasylirii + pollen (33.7 eggs/female), which was statistically different from those fed with F. dasylirii + honey (50.9 eggs/female) and with F. dasylirii alone (51.8 eggs/female), which, in turn, did not differ from each other ( F = 0.77; p = 0.46). In line with the results of the present study, Lima et al. (2020) found that adult females of another ladybug species, Brumoides foudrasii (Mulsant, 1850) (Coleoptera: Coccinellidae), did not oviposit when fed with F. dasylirii + pollen, and when fed with pollen alone, there was no oviposition or mature oocytes, indicating that pollen inhibited oviposition in the mentioned species. Guerreiro et al. (2003) also did not find significant differences in the fecundity of females of another ladybug species, Pentilia egena Mulsant, 1850 (Coleoptera: Coccinellidae), when fed only its natural diet, the scale insect Aspidiotus nerii Bouché, 1833 (Hemiptera: Diaspididae), and when the natural diet was supplemented with honey or honey + yeast. Hodek and Honek (1996) report that these alternative foods would serve only as an energy source, not influencing fecundity, and Lundgren (2009) states that sugar is a poor source for reproduction. Taken together, the results of the present study demonstrate that supplementing the essential diet with alternative foods, such as honey and pollen, did not positively interfere with the reproductive performance of T. notata females, thereby not supporting the hypothesis that such foods would improve the predator's performance. Declarations Acknowledgements The authors are grateful to the Laboratory of Biological Control and Insect Ecology at UFRPE (Recife, PE, Brazil) for providing the insects used in the experiment. Author Contribution MSL and RB conceived and designed the experiment; MSL and BSM performed the experiment; ECG analyzed the data; BSM drafted the manuscript; WJTP, RB, and ECG reviewed and translated the work; RB acquired funding; all authors read and approved the final version of the manuscript. Funding This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001. Data Availability Data will be available from the corresponding author upon reasonable request. Ethical Approval Not applicable. Competing Interest The authors declare no competing interests. References Almeida LM, Ribeiro-Costa CS (2009) Coleópteros predadores (Coccinellidae). In: Panizzi AR, Parra JRP (ed) Bioecologia e nutrição de insetos: base para o manejo integrado de pragas. Embrapa Informação Tecnológica, Brasília, pp 931-968 Almeida LM, Ribeiro-Costa CS (2012) Predatory beetles (Coccinellidae). In: Panizzi AR, Parra JRP (ed) Insect bioecology and nutrition for integrated pest management. CRC Press, Boca Raton, pp 571-591 Berkvens N, Bonte J, Berkvens D, Deforce K, Tirry L, De Clercq P (2008) Pollen as an alternative food for Harmonia axyridis . BioControl 53(1):201-210. https://doi.org/10.1007/s10526-007-9128-7 Bertin A, Pinto AS, Parra JRP (2021) Controle de qualidade de inimigos naturais. In: Parra JRP, Pinto AS, Nava DE, Oliveira RC, Diniz AJF (ed) Controle biológico com parasitoides e predadores na agricultura brasileira. FEALQ, Piracicaba, pp 401-426 Bueno VHP, van Lenteren JC (2021) Insetos predadores no controle biológico aumentativo de pragas. In: Parra JRP, Pinto AS, Nava DE, Oliveira RC, Diniz AJF (ed) Controle biológico com parasitoides e predadores na agricultura brasileira. FEALQ, Piracicaba, pp 249-282 Cividanes TMS, Cividanes FJ, Matos BA (2007) Biologia de Psyllobora confluens alimentada com o fungo Erysiphe cichoracearum . Pesq Agropec Bras 42(12):1675-1679. https://doi.org/10.1590/S0100-204X2007001200002 Correia ACB, Berti Filho E (1988) Aspectos biológicos de Cycloneda zischkai Mader, 1950 (Coleoptera: Coccinellidae), predador de psilídeos. An Soc Entomol Bras 17(2):333-345. https://doi.org/10.37486/0301-8059.v17i2.526 De Clercq P, Bonte M, Van Speybroeck K, Bolckmans K, Deforce K (2005) Development and reproduction of Adalia bipunctulata (Coleoptera: Coccinellidae) on eggs of Ephestia kuehniella (Lepidoptera: Phycitidae) and pollen. Pest Manag Sci 61(11):1129-1132. https://doi.org/10.1002/ps.1111 Guerreiro JC, Berti Filho E, Busoli AC (2003) Influência da alimentação complementar na oviposição e eficiência de predação de Pentilia egena (Coleoptera: Coccinellidae) sobre Aspidiotus nerii (Hemiptera: Diaspididae). Rev Cient Eletron Agron 4:1-4 Hemptinne JL, Naisse J (1987) Ecophysiology of the reproductive activity of Adalia bipunctata L. (Coleoptera: Coccinellidae). Meded Fac Landbouww Rijksuniv Gent52:225-233 Herren HR, Neuenschwander P (1991) Biological control of cassava pests in Africa. Annu Rev Entomol 36(1):257-283. https://doi.org/10.1146/annurev.en.36.010191.001353 Hodek I (1973) Biology of Coccinellidae. Academy of Sciences, Prague Hodek I, Honek A (1996) Ecology of Coccinellidae. Kluwer Academic, Dordrecht Iperti G (1999) Biodiversity of predaceous Coccinellidae in relation to bioindication and economic importance. Agric Ecosyst Environ 74(1-3):323-342. https://doi.org/10.1016/S0167-8809(99)00041-9 Lima MS, Melo JWS, Barros R (2016) Biology of Zagreus bimaculosus Mulsant (Coleoptera: Coccinellidae), a predator of Ferrisia dasylirii (Cockerell) (Hemiptera: Pseudococcidae). Coleopt Bull 70(2):314-320. https://doi.org/10.1649/0010-065X-70.2.314 Lima MS, Melo JWS, Barros R (2018) Alternative food sources for the ladybird Brumoides foudrasii (Mulsant) (Coleoptera: Coccinellidae). Braz J Biol 78(2):211-216. https://doi.org/10.1590/1519-6984.02816 Lima MS, Silva DMP, Falcão HM, Ferreira WM, Silva LD, Paranhos BAJ (2011) Predadores associados á Dactylopius opuntiae (Hemiptera: Dactylopiidae) em palma forrageira no Estado de Pernambuco, Brasil. Rev Chilena Ent 36:51-54 Lima MS, Pontes WJT, Nóbrega RL (2020) Pollen did not provide suitable nutrients for ovary development in a ladybird Brumoides foudrasii (Coleoptera: Coccinellidae). Diversitas J 5(3):1486-1494. https://doi.org/10.17648/diversitas-journal-v5i3-877 Lundgren JG (2009) Nutritional aspects of non-prey foods in the life histories of predaceous Coccinellidae. Biol Control 51(2):294-305. https://doi.org/10.1016/j.biocontrol.2009.05.016 Neuenschwander P, Zweigert M (1994) Biological control of cassava mealybug and green spider mite in eastern and Southern Africa. In: Allard GB, Skoglund LG, Neuenschwander P, Murphy RJ (ed) Root and tuber crops pest management in East and Southern Africa: proceedings of a Workshop held in Mombasa, 1992. CABI, Nairobi, pp 13-18 Obrycki JJ, Kring TJ (1998) Predaceous Coccinellidae in biological control. Annu Rev Entomol 43:295-321. https://doi.org/10.1146/annurev.ento.43.1.295 Parra JRP (2021) Criação massal de inimigos naturais. In: Parra JRP, Pinto AS, Nava DE, Oliveira RC, Diniz AJF (ed) Controle biológico com parasitoides e predadores na agricultura brasileira. FEALQ, Piracicaba, pp 379-400 Parajulee MN, Phillips TW (1993) Effects of prey species on development and reproduction of the predator Lyctocoris campestris (Heteroptera: Anthocoridae). Environ Entomol 22(5):1035-1042. https://doi.org/10.1093/ee/22.5.1035 Pemberton RW, Vandenberg NJ (1993) Extrafloral nectar feeding by ladybird beetles (Coleoptera: Coccinellidae). Proc Entomol Soc Washington 95(2):139-151. SAS Institute (2001) SAS/STAT User’s guide, version 8.02, TS level 2MO. SAS Institute Inc., Cary Smith SF, Krischik VA (1999) Effects of systemic imidacloprid on Coleomegilla maculata (Coleoptera: Coccinellidae). Environ Entomol 28(6):1189-1195. https://doi.org/10.1093/ee/28.6.1189 Souza B, Santos-Cividanes TM, Cividanes FJ, Sousa ALV (2019) Predatory insects. In: Souza B, Vázquez LL, Marucci RC (ed) Natural enemies of insect pests in neotropical agroecosystems: biological control and functional biodiversity. Springer, Cham, pp 73-87 Souza B, Souza-Bezerra CE (2019) Predatory insects. In: Souza B, Vázquez LL, Marucci RC (ed) Natural enemies of insect pests in neotropical agroecosystems: biological control and functional biodiversity. Springer, Cham, pp 175-187 Stäubli Dreyer BS, Neuenschwander P, Baumgärtner J, Dorn, S (1997) Trophic influences on survival, development and reproduction of Hyperaspis notata (Col., Coccinellidae). J Appl Entomol 121:249-256. https://doi.org/10.1111/j.1439-0418.1997.tb01401.x Sujii ER, Pires CSS, Venzon M, Fernandes OA (2020) Controle de artrópodes-praga com insetos predadores. In: Fontes EMG, Valadares-Inglis MC (ed) Controle biológico de pragas da agricultura. Embrapa, Brasília, pp 113-140 Zanuncio JC, Guedes RNC, Oliveira HN, Zanuncio TV (2002) Uma década de estudos com percevejos predadores: conquistas e desafios. In: Parra JRP, Botelho PSM, Corrêa-Ferreira BS, Bento JMS (ed) Controle biológico no Brasil: parasitóides e predadores. Manole, São Paulo, pp 495-510 Table 1 Table 1 Reproductive parameters (mean ± SE) of adult females of Tenuisvalvae notata fed on Ferrisia dasylirii and supplemented with pollen and honey Parameters Food F. dasylirii F. dasylirii + pollen F. dasylirii + honey Pre-oviposition (days) 5.4 ± 0.41 a 5.2 ± 0.59 a 5.0 ± 0.44 a Oviposition (days) 15.7 ± 2.36 a 8.8 ± 1.60 a 12.4 ± 2.67 a Pos-oviposition (days) 16.4 ± 3.47 a 21.9 ± 6.02 a 16.6 ± 6.12 a Longevity (days) 36.2 ± 4.66 a 38.0 ± 5.79 a 32.3 ± 6.6 a Fecundity (eggs/female) 51.8 ± 1.80 a 33.7 ± 7.65 b 50.9 ±1.46 a Fertility (%) 75.3 ±1.99 a 71.4 ± 4.5 a 73.7 ± 3.14 a *Means followed by the same letters within a row do not differ (t-test: α = 0.05). Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 28 Oct, 2025 Read the published version in International Journal of Tropical Insect Science → Version 1 posted Editorial decision: Revision requested 21 Aug, 2025 Reviews received at journal 19 Aug, 2025 Reviews received at journal 18 Aug, 2025 Reviewers agreed at journal 13 Aug, 2025 Reviewers agreed at journal 11 Aug, 2025 Reviewers agreed at journal 08 Aug, 2025 Reviewers agreed at journal 08 Aug, 2025 Reviewers agreed at journal 08 Aug, 2025 Reviewers agreed at journal 07 Aug, 2025 Reviewers agreed at journal 06 Aug, 2025 Reviewers agreed at journal 06 Aug, 2025 Reviewers invited by journal 06 Aug, 2025 Editor assigned by journal 22 Jul, 2025 Submission checks completed at journal 22 Jul, 2025 First submitted to journal 18 Jul, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7160590","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Short Report","associatedPublications":[],"authors":[{"id":497940051,"identity":"f94741e8-f98d-492e-8e86-7c67fe714e05","order_by":0,"name":"Maurício Silva Lima","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABCElEQVRIiWNgGAWjYLCCBAYJBgZ2xgYGhgqoSAWDBYMBQS3MIC1noCJngCJ4tYABMxAzthGhRbf9jNmHBzUWcgzMzK0bfs6zk+NvP8AmcaBGgsFc+gBWLWZncoxnJByTMAY6rO1m77ZkY4kzCUAtxyQYLPsSsGs5kGPMkNggkdgA1HKDd9uBxA0MCWzSH9iADjuDVQeD2fk3CC03/845UL+B/wHQln94tNxAsuU2b8OBBAMJoMMOtuHT8qyYAeQXNpAWmWPJhjNuPGy2ONgnwWPZg8thyZsZf9TUyfGztz+7+abGTp6/P/ngjQPfbOTMebBrgQM2BBMUpwwMhDSMglEwCkbBKMADAPHtV3zSGe9/AAAAAElFTkSuQmCC","orcid":"","institution":"Universidade Federal de Alagoas – UFAL","correspondingAuthor":true,"prefix":"","firstName":"Maurício","middleName":"Silva","lastName":"Lima","suffix":""},{"id":497940052,"identity":"bf5f8daf-5818-4340-b303-d3c7f1f3caaa","order_by":1,"name":"Wendel José Teles Pontes","email":"","orcid":"","institution":"Universidade Federal de Pernambuco – UFPE","correspondingAuthor":false,"prefix":"","firstName":"Wendel","middleName":"José Teles","lastName":"Pontes","suffix":""},{"id":497940053,"identity":"48acba36-9c7b-4637-b542-be983d57ceba","order_by":2,"name":"Elio Cesar Guzzo","email":"","orcid":"","institution":"Unidade de Execução de Pesquisa de Rio Largo","correspondingAuthor":false,"prefix":"","firstName":"Elio","middleName":"Cesar","lastName":"Guzzo","suffix":""},{"id":497940054,"identity":"e3f1fe10-8e1e-4d2e-982b-637d877c6309","order_by":3,"name":"Bruno Silva Monteiro","email":"","orcid":"","institution":"Universidade Federal Rural de Pernambuco – UFRPE","correspondingAuthor":false,"prefix":"","firstName":"Bruno","middleName":"Silva","lastName":"Monteiro","suffix":""},{"id":497940055,"identity":"c2c20781-71d2-4aca-a4b0-bfb64107a6eb","order_by":4,"name":"Reginaldo Barros","email":"","orcid":"","institution":"Universidade Federal Rural de Pernambuco – UFRPE","correspondingAuthor":false,"prefix":"","firstName":"Reginaldo","middleName":"","lastName":"Barros","suffix":""}],"badges":[],"createdAt":"2025-07-18 20:53:10","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7160590/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7160590/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s42690-025-01653-x","type":"published","date":"2025-10-28T15:57:16+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":95039904,"identity":"714b41ce-a579-45b1-b9d8-de8b067251d7","added_by":"auto","created_at":"2025-11-03 16:05:19","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":409095,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7160590/v1/1a97977c-0d31-4d71-9248-3622308f99fe.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"\u003cp\u003eDo honey and pollen enhance the reproductive performance of ladybugs? A case study on Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae)\u003c/p\u003e","fulltext":[{"header":"Full Text","content":"\u003cp\u003eLadybugs, also known as ladybird beetles (Coleoptera: Coccinellidae), are among the most important predators of pests in agricultural crops (Obrycki and Kring 1998; Iperti 1999; Souza et al. 2019; Sujii et al. 2020; Bueno and van Lenteren 2021). These beetles are the primary predators of scale insects (Hemiptera: Diaspididae, Dactylopiidae, and Pseudococcidae) (Lima et al. 2011, 2016, 2018), with which they are often found associated, and are considered their essential food source. The ladybug \u003cem\u003eTenuisvalvae notata\u003c/em\u003e (Mulsant, 1850) (Coleoptera: Coccinellidae), in both larval and adult stages, preys on scale insects and is used, for example, as a complement to biological control carried out by the parasitoid \u003cem\u003eAnagyrus lopezi\u003c/em\u003e (De Santis, 1964) (Hymenoptera: Encyrtidae) on the mealybug \u003cem\u003ePhenacoccus manihoti\u003c/em\u003e Matile-Ferrero, 1977 (Hemiptera: Pseudococcidae) in cassava \u003cem\u003eManihot esculenta\u003c/em\u003e Crantz crops in Africa (St\u0026auml;ubli Dreyer et al. 1997).\u003c/p\u003e\n\u003cp\u003eThe efficient mass production of natural enemies is a fundamental tool within Integrated Pest Management, aiming to provide parasitoids and predators with biological and behavioral characteristics compatible with natural populations and capable of reducing the population density of economically important pests while maintaining economic viability in the production system (Zanuncio et al. 2002; Souza and Souza-Bezerra 2019; Bertin et al. 2021; Parra 2021). To achieve this, predatory insects rely on factors such as the quality and quantity of food to mitigate adverse effects on their biological aspects, such as development, reproduction, and longevity (Parajulee and Phillips 1993).\u003c/p\u003e\n\u003cp\u003eAlternative foods, such as pollen, nectar, honeydew, honey, fruits, foliage, and fungi (Pemberton and Vandenberg 1993; Cividanes et al. 2007; Lundgren 2009), can help to maintain predator populations in their habitat in response to changes in pest population density (Lundgren 2009). Although pollen does not play a vital role in the ladybugs\u0026apos; diet, 39 species of ladybugs that consume pollen from 88 plant species have been identified, under both field and laboratory conditions (Lundgren 2009). Pollen can promote spermatogenesis (Hemptinne and Naisse 1987) and, when combined with prey, can aid in the reproduction of certain species (De Clercq et al. 2005; Berkvens et al. 2008). Carbohydrate-rich foods, such as honey, which is rich in glucose and fructose, can significantly improve the adult survival of ladybirds and, when combined with essential food, can allow females to enhance their reproductive performance (Lundgren 2009).\u003c/p\u003e\n\u003cp\u003eEssential foods support the development and reproduction of ladybugs, while alternative foods are helpful as an energy source, prolonging their survival (Hodek and Honek 1996; Almeida and Ribeiro-Costa 2009, 2012). Many studies have proposed that essential food, combined with alternative food, can enhance the performance of ladybugs, serving as a supplement in prey scarcity, as they actively seek these alternative foods even when prey is abundant. Therefore, the simultaneous consumption of both types of food would increase the performance of these predators (Lundgren 2009). However, other studies suggest that dietary supplementation with pollen may have a negative influence on ladybug reproduction (Lima et al. 2020).\u003c/p\u003e\n\u003cp\u003eThus, the present study aimed to evaluate the effect of dietary supplementation on the biological performance of \u003cem\u003eT. notata\u003c/em\u003e females, testing the hypothesis that supplementary food should optimize the biological parameters of this species when compared to individuals exclusively fed on their essential food, the mealybug \u003cem\u003eFerrisia dasylirii\u003c/em\u003e (Cockerell, 1896) (Hemiptera: Pseudococcidae), under laboratory conditions.\u003c/p\u003e\n\u003cp\u003eThe experiment was conducted at the Laboratory of Plant Biology and Insect Resistance, located in the Department of Agronomy at the Federal Rural University of Pernambuco (UFRPE), in Recife, Pernambuco, Brazil. Individuals of \u003cem\u003eF. dasylirii\u003c/em\u003e and \u003cem\u003eT. notata\u003c/em\u003e used in the experiments were obtained from the insect rearing facility provided by the Laboratory of Biological Control and Insect Ecology at UFRPE. The rearing facilities were established in a climate-controlled room with controlled conditions (25 \u0026plusmn; 2 \u0026deg;C, 70 \u0026plusmn; 10% relative humidity, and a 12-hour photoperiod).\u003c/p\u003e\n\u003cp\u003eTo rear \u003cem\u003eF. dasylirii\u003c/em\u003e, pumpkins \u003cem\u003eCucurbita maxima\u003c/em\u003e Duch (Cucurbitaceae) cv. Jacarezinho, in its initial stage of ripening, was used as a host plant. All fruits were sanitized with a 10% sodium hypochlorite solution before infestation. The pumpkins were placed on a glass support inside plastic trays (30 \u0026times; 45 cm), which were in turn placed on a larger plastic tray (64 \u0026times; 63 cm) containing water + detergent to prevent infestation by ants and other undesirable insects. Infested fruits were placed alongside non-infested fruits, allowing nymphs to move from one to the other. In the absence of nymphs, females were transferred to non-infested fruits using a fine-bristle brush.\u003c/p\u003e\n\u003cp\u003ePlastic cages (40 \u0026times; 25 \u0026times; 20 cm) covered with voile were used for rearing \u003cem\u003eT. notata\u003c/em\u003e. Infested pumpkins with \u003cem\u003eF. dasylirii\u003c/em\u003e were provided as food. To prevent damage to the fruit and its rapid deterioration, the pumpkins were placed on a metal support protected by a sponge. As a supplementary food source, honeybee pollen, 5% honey, or eggs of \u003cem\u003eEphestia kuehniella\u003c/em\u003e Zeller, 1879 (Lepidoptera: Pyralidae) were offered. Young and adult forms were reared separately. Pumpkins were periodically replaced with newly infested fruits or when they became deteriorated.\u003c/p\u003e\n\u003cp\u003eTo obtain \u003cem\u003eT. notata\u003c/em\u003e individuals of known and standardized age, pairs of the insect from the rearing facility were separated into plastic containers (300 mL) and fed with \u003cem\u003eF. dasylirii\u003c/em\u003e adults. Egg masses were removed after 24 hours, and the larvae from these eggs were fed with \u003cem\u003eF. dasylirii\u003c/em\u003e nymphs until the emergence of adults, as preliminary tests showed that the larvae are highly specific and do not feed on pollen and honey. Pairs of \u003cem\u003eT. notata\u003c/em\u003e adults of the same age were removed from the rearing cage and individually placed in plastic containers (300 mL) containing one of the following treatments. The containers were labeled and kept in a B.O.D. (Biological Oxygen Demand) incubator at a temperature of 25 \u0026plusmn; 1 \u0026deg;C, relative humidity of 70 \u0026plusmn; 10%, and a 12-hour photophase. Male \u003cem\u003eT. notata\u003c/em\u003e were removed from the containers after 72 hours.\u003c/p\u003e\n\u003cp\u003eThe experiment consisted of three treatments: 1) females fed only with \u003cem\u003eF. dasylirii\u003c/em\u003e; 2) females fed with \u003cem\u003eF. dasylirii\u003c/em\u003e + honey; and 3) females fed with \u003cem\u003eF. dasylirii\u003c/em\u003e + pollen. Pure honey (diluted to 5% for the experiments) and commercial honeybee pollen were used. Mealybugs were offered daily \u003cem\u003ead libitum\u003c/em\u003e directly in the container, and honey and pollen were offered, respectively, on moistened cotton and in small plastic containers, and replaced every 3 days until the death of the females. The investigated biological parameters included the pre-oviposition, oviposition, and post-oviposition periods, as well as longevity, fertility, and fecundity. The experimental design was completely randomized, with 20 replications, where each container with a \u003cem\u003eT. notata\u003c/em\u003e female was considered as one replication. Means were subjected to analysis of variance and compared using the t-test at a 5% significance level, using the PROC ANOVA function of SAS statistical software (SAS Institute 2001).\u003c/p\u003e\n\u003cp\u003eThe results showed that feeding adult female \u003cem\u003eT. notata\u003c/em\u003e with nymphs of \u003cem\u003eF. dasylirii\u003c/em\u003e, nymphs of \u003cem\u003eF. dasylirii\u003c/em\u003e + pollen, or nymphs of \u003cem\u003eF. dasylirii\u003c/em\u003e + honey did not cause significant differences in the pre-oviposition (5.4, 5.2, and 5.0 days, respectively) (\u003cem\u003eF\u003c/em\u003e = 0.11; \u003cem\u003ep\u003c/em\u003e = 0.89), oviposition (15.7, 8.8, and 15.7 days, respectively) (\u003cem\u003eF\u003c/em\u003e = 2.43; \u003cem\u003ep\u003c/em\u003e = 0.09), or post-oviposition (16.4, 21.9, and 16.6 days, respectively) (\u003cem\u003eF\u003c/em\u003e = 0.40; \u003cem\u003ep\u003c/em\u003e = 0.67) periods, which was reflected in their longevity (36.2, 38.0, and 32.3 days, respectively) (\u003cem\u003eF\u003c/em\u003e = 0.18; p = 0.83) (Table 1), which encompasses the pre-oviposition, oviposition, and post-oviposition periods combined.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 1 here\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eCorreia and Berti Filho (1988) highlight that complementary feeding serves as an energy source that can only prolong the survival of adult ladybugs. Additionally, Smith and Krischik (1999) suggest that sugar-based feeding can reduce the pre-oviposition period of Coccinellidae and help females to survive during diapause periods. In Africa, pure honey serves as a food source for adult \u003cem\u003eT. notata\u003c/em\u003e, which is employed in the biological control of \u003cem\u003eP. manihoti\u003c/em\u003e in cassava, allowing ladybugs to maintain their longevity during transportation (Herren and Neuenschwander 1991; Neuenschwander and Zweigert 1994). However, such effects were not observed in the present study and appear not to apply to \u003cem\u003eT. notata\u003c/em\u003e and the provided foods. As Lundgren (2009) suggests, alternative food serves to stabilize the predator population during periods of prey scarcity when it diversifies its food type in response to changes in the population density of its essential food source within its niche. However, in the current study, there was no restriction on the natural prey (\u003cem\u003eF. dasylirii\u003c/em\u003e) for the predator \u003cem\u003eT. notata\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eRegarding fecundity, \u003cem\u003eT. notata\u0026nbsp;\u003c/em\u003efemales laid fewer eggs when fed with \u003cem\u003eF. dasylirii\u003c/em\u003e + pollen (33.7 eggs/female), which was statistically different from those fed with \u003cem\u003eF. dasylirii\u003c/em\u003e + honey (50.9 eggs/female) and with \u003cem\u003eF. dasylirii\u003c/em\u003e alone (51.8 eggs/female), which, in turn, did not differ from each other (\u003cem\u003eF\u003c/em\u003e = 0.77; \u003cem\u003ep\u003c/em\u003e = 0.46). In line with the results of the present study, Lima et al. (2020) found that adult females of another ladybug species, \u003cem\u003eBrumoides foudrasii\u003c/em\u003e (Mulsant, 1850) (Coleoptera: Coccinellidae), did not oviposit when fed with \u003cem\u003eF. dasylirii\u003c/em\u003e + pollen, and when fed with pollen alone, there was no oviposition or mature oocytes, indicating that pollen inhibited oviposition in the mentioned species.\u003c/p\u003e\n\u003cp\u003eGuerreiro et al. (2003) also did not find significant differences in the fecundity of females of another ladybug species, \u003cem\u003ePentilia egena\u003c/em\u003e Mulsant, 1850 (Coleoptera: Coccinellidae), when fed only its natural diet, the scale insect \u003cem\u003eAspidiotus nerii\u003c/em\u003e Bouch\u0026eacute;, 1833 (Hemiptera: Diaspididae), and when the natural diet was supplemented with honey or honey + yeast. Hodek and Honek (1996) report that these alternative foods would serve only as an energy source, not influencing fecundity, and Lundgren (2009) states that sugar is a poor source for reproduction.\u003c/p\u003e\n\u003cp\u003eTaken together, the results of the present study demonstrate that supplementing the essential diet with alternative foods, such as honey and pollen, did not positively interfere with the reproductive performance of \u003cem\u003eT. notata\u003c/em\u003e females, thereby not supporting the hypothesis that such foods would improve the predator\u0026apos;s performance.\u003c/p\u003e\n"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors are grateful to the Laboratory of Biological Control and Insect Ecology at UFRPE (Recife, PE, Brazil) for providing the insects used in the experiment.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAuthor Contribution\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eMSL and RB conceived and designed the experiment; MSL and BSM performed the experiment; ECG analyzed the data; BSM drafted the manuscript; WJTP, RB, and ECG reviewed and translated the work; RB acquired funding; all authors read and approved the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was financed in part by the Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior - Brasil (CAPES) - Finance Code 001.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eData Availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData will be available from the corresponding author upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical Approval\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eCompeting Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlmeida LM, Ribeiro-Costa CS (2009) Cole\u0026oacute;pteros predadores (Coccinellidae). In: Panizzi AR, Parra JRP (ed) Bioecologia e nutri\u0026ccedil;\u0026atilde;o de insetos: base para o manejo integrado de pragas. Embrapa Informa\u0026ccedil;\u0026atilde;o Tecnol\u0026oacute;gica, Bras\u0026iacute;lia, pp 931-968\u003c/li\u003e\n\u003cli\u003eAlmeida LM, Ribeiro-Costa CS (2012) Predatory beetles (Coccinellidae). In: Panizzi AR, Parra JRP (ed) Insect bioecology and nutrition for integrated pest management. CRC Press, Boca Raton, pp 571-591\u003c/li\u003e\n\u003cli\u003eBerkvens N, Bonte J, Berkvens D, Deforce K, Tirry L, De Clercq P (2008) Pollen as an alternative food for \u003cem\u003eHarmonia axyridis\u003c/em\u003e. BioControl 53(1):201-210. https://doi.org/10.1007/s10526-007-9128-7\u003c/li\u003e\n\u003cli\u003eBertin A, Pinto AS, Parra JRP (2021) Controle de qualidade de inimigos naturais. In: Parra JRP, Pinto AS, Nava DE, Oliveira RC, Diniz AJF (ed) Controle biol\u0026oacute;gico com parasitoides e predadores na agricultura brasileira. 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Agric Ecosyst Environ 74(1-3):323-342. https://doi.org/10.1016/S0167-8809(99)00041-9\u003c/li\u003e\n\u003cli\u003eLima MS, Melo JWS, Barros R (2016) Biology of \u003cem\u003eZagreus bimaculosus\u003c/em\u003e Mulsant (Coleoptera: Coccinellidae), a predator of \u003cem\u003eFerrisia dasylirii\u003c/em\u003e (Cockerell) (Hemiptera: Pseudococcidae). Coleopt Bull 70(2):314-320. https://doi.org/10.1649/0010-065X-70.2.314\u003c/li\u003e\n\u003cli\u003eLima MS, Melo JWS, Barros R (2018) Alternative food sources for the ladybird \u003cem\u003eBrumoides foudrasii\u003c/em\u003e (Mulsant) (Coleoptera: Coccinellidae). Braz J Biol 78(2):211-216. https://doi.org/10.1590/1519-6984.02816\u003c/li\u003e\n\u003cli\u003eLima MS, Silva DMP, Falc\u0026atilde;o HM, Ferreira WM, Silva LD, Paranhos BAJ (2011) Predadores associados \u0026aacute; \u003cem\u003eDactylopius opuntiae\u003c/em\u003e (Hemiptera: Dactylopiidae) em palma forrageira no Estado de Pernambuco, Brasil. Rev Chilena Ent 36:51-54\u003c/li\u003e\n\u003cli\u003eLima MS, Pontes WJT, N\u0026oacute;brega RL (2020) Pollen did not provide suitable nutrients for ovary development in a ladybird \u003cem\u003eBrumoides foudrasii\u003c/em\u003e (Coleoptera: Coccinellidae). Diversitas J 5(3):1486-1494. https://doi.org/10.17648/diversitas-journal-v5i3-877\u003c/li\u003e\n\u003cli\u003eLundgren JG (2009) Nutritional aspects of non-prey foods in the life histories of predaceous Coccinellidae. Biol Control 51(2):294-305. https://doi.org/10.1016/j.biocontrol.2009.05.016\u003c/li\u003e\n\u003cli\u003eNeuenschwander P, Zweigert M (1994) Biological control of cassava mealybug and green spider mite in eastern and Southern Africa. In: Allard GB, Skoglund LG, Neuenschwander P, Murphy RJ (ed) Root and tuber crops pest management in East and Southern Africa: proceedings of a Workshop held in Mombasa, 1992. CABI, Nairobi, pp 13-18\u003c/li\u003e\n\u003cli\u003eObrycki JJ, Kring TJ (1998) Predaceous Coccinellidae in biological control. Annu Rev Entomol 43:295-321. https://doi.org/10.1146/annurev.ento.43.1.295\u003c/li\u003e\n\u003cli\u003eParra JRP (2021) Cria\u0026ccedil;\u0026atilde;o massal de inimigos naturais. In: Parra JRP, Pinto AS, Nava DE, Oliveira RC, Diniz AJF (ed) Controle biol\u0026oacute;gico com parasitoides e predadores na agricultura brasileira. FEALQ, Piracicaba, pp 379-400\u003c/li\u003e\n\u003cli\u003eParajulee MN, Phillips TW (1993) Effects of prey species on development and reproduction of the predator \u003cem\u003eLyctocoris campestris\u003c/em\u003e (Heteroptera: Anthocoridae). Environ Entomol 22(5):1035-1042. https://doi.org/10.1093/ee/22.5.1035\u003c/li\u003e\n\u003cli\u003ePemberton RW, Vandenberg NJ (1993) Extrafloral nectar feeding by ladybird beetles (Coleoptera: Coccinellidae). Proc Entomol Soc Washington 95(2):139-151.\u003c/li\u003e\n\u003cli\u003eSAS Institute (2001) SAS/STAT User\u0026rsquo;s guide, version 8.02, TS level 2MO. SAS Institute Inc., Cary\u003c/li\u003e\n\u003cli\u003eSmith SF, Krischik VA (1999) Effects of systemic imidacloprid on \u003cem\u003eColeomegilla maculata\u003c/em\u003e (Coleoptera: Coccinellidae). Environ Entomol 28(6):1189-1195. https://doi.org/10.1093/ee/28.6.1189\u003c/li\u003e\n\u003cli\u003eSouza B, Santos-Cividanes TM, Cividanes FJ, Sousa ALV (2019) Predatory insects. In: Souza B, V\u0026aacute;zquez LL, Marucci RC (ed) Natural enemies of insect pests in neotropical agroecosystems: biological control and functional biodiversity. Springer, Cham, pp 73-87\u003c/li\u003e\n\u003cli\u003eSouza B, Souza-Bezerra CE (2019) Predatory insects. In: Souza B, V\u0026aacute;zquez LL, Marucci RC (ed) Natural enemies of insect pests in neotropical agroecosystems: biological control and functional biodiversity. Springer, Cham, pp 175-187\u003c/li\u003e\n\u003cli\u003eSt\u0026auml;ubli Dreyer BS, Neuenschwander P, Baumg\u0026auml;rtner J, Dorn, S (1997) Trophic influences on survival, development and reproduction of \u003cem\u003eHyperaspis notata\u003c/em\u003e (Col., Coccinellidae). J Appl Entomol 121:249-256. https://doi.org/10.1111/j.1439-0418.1997.tb01401.x\u003c/li\u003e\n\u003cli\u003eSujii ER, Pires CSS, Venzon M, Fernandes OA (2020) Controle de artr\u0026oacute;podes-praga com insetos predadores. In: Fontes EMG, Valadares-Inglis MC (ed) Controle biol\u0026oacute;gico de pragas da agricultura. Embrapa, Bras\u0026iacute;lia, pp 113-140\u003c/li\u003e\n\u003cli\u003eZanuncio JC, Guedes RNC, Oliveira HN, Zanuncio TV (2002) Uma d\u0026eacute;cada de estudos com percevejos predadores: conquistas e desafios. In: Parra JRP, Botelho PSM, Corr\u0026ecirc;a-Ferreira BS, Bento JMS (ed) Controle biol\u0026oacute;gico no Brasil: parasit\u0026oacute;ides e predadores. Manole, S\u0026atilde;o Paulo, pp 495-510\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table 1","content":"\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e Reproductive parameters (mean \u0026plusmn; SE) of adult females of \u003cem\u003eTenuisvalvae notata\u003c/em\u003e fed on \u003cem\u003eFerrisia dasylirii\u003c/em\u003e and supplemented with pollen and honey\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"489\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 149px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eParameters\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" style=\"width: 340px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eFood\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eF. dasylirii\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eF. dasylirii\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;+ pollen\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eF. dasylirii\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;+ honey\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003ePre-oviposition (days)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e5.4 \u0026plusmn; 0.41 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e5.2 \u0026plusmn; 0.59 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e5.0 \u0026plusmn; 0.44 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003eOviposition (days)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e15.7 \u0026plusmn; 2.36 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e8.8 \u0026plusmn; 1.60 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e12.4 \u0026plusmn; 2.67 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003e\u0026nbsp;Pos-oviposition (days)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e16.4 \u0026plusmn; 3.47 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e21.9 \u0026plusmn; 6.02 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e16.6 \u0026plusmn; 6.12 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003eLongevity (days)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e36.2 \u0026plusmn; 4.66 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e38.0 \u0026plusmn; 5.79 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e32.3 \u0026plusmn; 6.6 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003eFecundity (eggs/female)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e51.8 \u0026plusmn; 1.80 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e33.7 \u0026plusmn; 7.65 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e50.9 \u0026plusmn;1.46 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 149px;\"\u003e\n \u003cp\u003eFertility (%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 85px;\"\u003e\n \u003cp\u003e75.3 \u0026plusmn;1.99 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 132px;\"\u003e\n \u003cp\u003e71.4 \u0026plusmn; 4.5 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003e73.7 \u0026plusmn; 3.14 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e*Means followed by the same letters within a row do not differ (t-test: \u0026alpha; = 0.05).\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Nutricional ecology, Insect nutrition, Insect feeding, Reproductive fitness","lastPublishedDoi":"10.21203/rs.3.rs-7160590/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7160590/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eCoccinellids (Coleoptera: Coccinellidae) are omnivorous insects. In addition to their primary prey, which is the essential food, they also consume alternative foods such as pollen, honey, and fungi. Recent studies have suggested that the combination of essential and alternative foods optimizes the performance of coccinellids, with the alternative food acting as a supplement, in the absence of the essential one. The alternative food would be actively sought, even if the essential one was abundant, as the simultaneous consumption of both would improve the predator's performance. \u003cem\u003eTenuisvalvae notata\u003c/em\u003e (Mulsant, 1850) (Coleoptera: Coccinellidae) is an important predator, primarily of mealybugs. To test the hypothesis, we carried out an experiment that consisted of three treatments: adult females of \u003cem\u003eT. notata\u003c/em\u003e fed only with the mealybug \u003cem\u003eFerrisia dasylirii\u003c/em\u003e (Cockerell, 1896) (Hemiptera: Pseudococcidae); with \u003cem\u003eF. dasylirii\u003c/em\u003e\u0026thinsp;+\u0026thinsp;pollen; or with \u003cem\u003eF. dasylirii\u003c/em\u003e\u0026thinsp;+\u0026thinsp;bee honey. There was no statistical difference among the three treatments regarding periods of pre-oviposition (5.4, 5.2, and 5.0 days, respectively), oviposition (15.7, 8.8, and 15.7 days, respectively), and post-oviposition (16.4, 21.9, and 16.6 days, respectively), longevity (36.2, 38.0, and 32.3 days, respectively), and fertility (75.3%, 71.4%, and 73.7%, respectively) of \u003cem\u003eT. notata\u003c/em\u003e females. Supplementation of the essential food with pollen significantly reduced fecundity (33.7 eggs/female) in relation to honey and the essential food alone (50.9 and 51.8 eggs/female, respectively), which, in turn, did not differ from each other. The results refuted the hypothesis that alternative food would optimize the reproductive performance of \u003cem\u003eT. notata\u003c/em\u003e females.\u003c/p\u003e","manuscriptTitle":"Do honey and pollen enhance the reproductive performance of ladybugs? A case study on Tenuisvalvae notata (Mulsant, 1850) (Coleoptera: Coccinellidae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-08-11 22:23:32","doi":"10.21203/rs.3.rs-7160590/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-08-21T06:39:08+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-19T12:03:31+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-18T06:29:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"260120432732594411757242223386630997298","date":"2025-08-13T05:20:47+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"259769700384631017621390274407600738106","date":"2025-08-11T21:12:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"39889393027069000403705733986296819031","date":"2025-08-09T03:53:08+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"233936640992449612050232443134837650168","date":"2025-08-09T01:50:56+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"65235404196592105039677962716749799881","date":"2025-08-08T16:12:52+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"97003022521184196455188292644743480650","date":"2025-08-07T05:25:20+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"104515454864916329210363736586576164507","date":"2025-08-07T03:07:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"141964606606072847616345624279706172164","date":"2025-08-06T20:57:49+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-08-06T18:15:28+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-07-22T12:53:38+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-07-22T12:52:08+00:00","index":"","fulltext":""},{"type":"submitted","content":"International Journal of Tropical Insect Science","date":"2025-07-18T20:43:31+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"8b59d198-3604-497f-8d8b-50ecc3ebc5ab","owner":[],"postedDate":"August 11th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-11-03T16:00:13+00:00","versionOfRecord":{"articleIdentity":"rs-7160590","link":"https://doi.org/10.1007/s42690-025-01653-x","journal":{"identity":"international-journal-of-tropical-insect-science","isVorOnly":false,"title":"International Journal of Tropical Insect Science"},"publishedOn":"2025-10-28 15:57:16","publishedOnDateReadable":"October 28th, 2025"},"versionCreatedAt":"2025-08-11 22:23:32","video":"","vorDoi":"10.1007/s42690-025-01653-x","vorDoiUrl":"https://doi.org/10.1007/s42690-025-01653-x","workflowStages":[]},"version":"v1","identity":"rs-7160590","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7160590","identity":"rs-7160590","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}