Ekolojik Zararlı Mücadelesinde Alternatif Stratejiler: Galleria mellonella ve Tenebrio molitor’a Karşı Entomopatojen Funguslar ile Bitkisel Uçucu Yağların Etkileşimi

Abstract

Beauveria bassiana ve Metarhizium brunneum gibi entomopatojen funguslar önemli biyolojik mücadele etmenleri arasında yer almakla birlikte, nispeten yavaş etki göstermeleri etkinliklerini sınırlayabilmektedir. Biyoaktif bileşenlerce zengin olan bitkisel uçucu yağlar, insektisit ve kovucu özellikleri nedeniyle potansiyel tamamlayıcı ajanlar olarak değerlendirilmektedir. Bu çalışmada, B. bassiana (BMAUM-M6-4) ve M. brunneum (V275) izolatlarının kekik, biberiye ve okaliptüs uçucu yağları ile Galleria mellonella ve Tenebrio molitor larvalarına karşı tekli ve kombine etkileri incelenmiştir. Uçucu yağlar tek başına, fungal süspansiyonlarla eş zamanlı veya 24 saat aralıkla uygulanarak etkileşimleri değerlendirilmiştir. Ayrıca, uçucu yağların fungusların miselyum gelişimi ve konidiyal çimlenmesi üzerindeki olası engelleyici etkilerini belirlemek için in-vitro testler gerçekleştirilmiştir. Sonuçlar, kekik yağının her iki fungus izolatı ile en güçlü sinerjistik etkiyi oluşturduğunu göstermiştir. Kekik yağı ile B. bassiana veya M. brunneum’un eş zamanlı uygulanması, tekli uygulamalara kıyasla larva mortalitesini anlamlı derecede artırmıştır. Biberiye yağı orta düzeyde uyumluluk göstermiş ancak larvaların biberiye ile ön işlem görmesi durumunda fungusun etkinliği azalmıştır. Okaliptüs yağı ise mortaliteye sınırlı katkı sağlamıştır. Önemli olarak, in-vitro testler uçucu yağların %1 konsantrasyonda fungal spor çimlenmesini anlamlı şekilde engellemediğini, miselyum gelişiminde ise yalnızca küçük düşüşler gözlendiğini ve bu etkiler arasında en düşük etkinin okaliptüs yağında ortaya çıktığını göstermiştir. Genel olarak, özellikle kekik ve biberiye yağlarının entomopatojen funguslarla kombinasyonu zararlı mortalitesini artırmış ve entegre zararlı yönetimi stratejileri kapsamında çevre dostu bir alternatif olarak potansiyel göstermiştir. Bununla birlikte, uygulama sırası ve konak türüne bağlı olarak etkinlik değişmekte olup, yarı-tarla ve tarla koşullarında optimizasyon ve doğrulamaya ihtiyaç duyulmaktadır.

Keywords

• Beauveria bassiana
• Metarhizium brunneum
• uçucu yağlar
• biyolojik mücadele
• etkileşim

Towards Eco-Friendly Pest Control: Synergistic Potential of Entomopathogenic Fungi and Plant Essential Oils Against Galleria mellonella and Tenebrio molitor

Abstract

Entomopathogenic fungi such as Beauveria bassiana and Metarhizium brunneum are important biological control agents, but their relatively slow action can limit efficiency. Essential oils, rich in bioactive compounds, represent potential complementary tools due to their insecticidal and repellent properties. The present study evaluated the individual and combined effects of B. bassiana (BMAUM-M6-4) and M. brunneum (V275) with thyme, rosemary, and eucalyptus essential oils against Galleria mellonella and Tenebrio molitor larvae. Essential oils were applied individually, simultaneously with fungal suspensions, or sequentially with a 24-hour interval to assess interactions. In addition, in vitro assays were conducted to determine the potential inhibitory effects of essential oils on fungal mycelial growth and conidial germination. The results revealed that thyme oil produced the strongest synergistic effect with both fungal isolates. Simultaneous application of thyme oil with B. bassiana or M. brunneum significantly increased larval mortality compared to single treatments. Rosemary oil showed moderate compatibility, although pre-exposure of larvae to rosemary reduced fungal efficacy. Eucalyptus oil contributed little to mortality under laboratory conditions. Importantly, in vitro assays showed that essential oils at 1% concentration did not significantly inhibit fungal spore germination, and only minor reductions in mycelial growth were observed, particularly with eucalyptus oil showing the least effect. Overall, combining entomopathogenic fungi with selected essential oils, particularly thyme and rosemary, enhanced pest mortality and demonstrated potential as an environmentally friendly alternative within integrated pest management strategies. Nevertheless, the effectiveness of these treatments depends on application sequence and host species, underscoring the need for optimization and validation under semi-field and field conditions.

Keywords

• Beauveria bassiana
• Metarhizium brunneum
• essential oils
• biological control
• interaction.

References

1. Abbott W.S., 1925. A method of computing the effectiveness of an insecticide. Journal of Economic Entomology, 18, 265-267.
2. Arnason J.T., Philogene B.J.R., Morand P., 1989. Insecticides of plant origin. ACS Symposium Series No 387, American Chemical Society, Washington DC, USA, 213 p.
3. Ayvaz A., Sagdic O., Karaborklu S., Ozturk I., 2010. Insecticidal activity of the essential oils from different plants against three stored product insects. Journal of Insect Science 10, 21. https://doi.org/10.1673/031.010.2101
4. Bardgett R., van der Putten W., 2014. Belowground biodiversity and ecosystem functioning. Nature, 515, 505–511. https://doi.org/10.1038/nature13855
5. Batish D.R., Singh H.P., Kohli R.K., Kaur S., 2008. Eucalyptus essential oil as natural pesticide. Forest Ecology and Management, 256 (12), 2166-2174. https://doi.org/10.1016/j.foreco.2008.08.008
6. Bedini S., Farina P., Napoli E., Flamini G., Ascrizzi R., Verzera A., Conti B., Zappalà L., 2021. Bioactivity of different chemotypes of oregano essential oil against the blowfly Calliphora vomitoria vector of foodborne pathogens. Insects, 12 (1), 52. https://doi.org/10.3390/insects12010052
7. Benelli G., Pavela R., Petrelli R., Cappellacci L., Bartolucci F., Canale A., Maggi F., 2019. Origanum syriacum subsp. syriacum: from an ingredient of Lebanese ‘manoushe’ to a source of effective and eco-friendly botanical insecticides. Industrial Crops and Products, 134, 26–32. https://doi.org/10.1016/j.indcrop.2019.03.055
8. Birah A., Chilana P., Shukla U.K., Gupta G.P., 2008. Mass rearing of greater wax moth (Galleria mellonella L.) on artificial diet. Indian Journal of Entomology, 70, 389–392.

9. Bitencourt R.O.B., Faria S.F., Marchesini P., dos Santos-Mallet J.R., Camargo M.G., Bittencourt V.R.E.P., Angelo I.C., 2022. Entomopathogenic fungi and Schinus molle essential oil: the combination of two eco-friendly agents against Aedes aegypti larvae. Journal of Invertebrate Pathology, 194, 107827. https://doi.org/10.1016/j.jip.2022.107827
10. Boucias D.G., Pendland J.C., 1982. Ultrastructural studies on the fungus, Nomuraea rileyi, infecting the velvetbean caterpillar, Anticarsia gemmatalis. Journal of Invertebrate Pathology, 39, 338-345.
11. Brooker M.I.H., Kleinig D.A., 2016. Field guide to Eucalypts. Volume 2. South-western and Southern Australia, Edition:3 pp. vii+-433.
12. Bruck D.J., 2005. Ecology of Metarhizium anisopliae in soilless potting media and the rhizosphere: Implications for pest management. Biological Control, 32 (1), 155–163. https://doi.org/10.1016/j.biocontrol.2004.09.003
13. Butt T.M., 2002. Use of entomogenous fungi for the control of insect pests. In: Agricultural applications. Esser K., Bennett JW (Eds.). The Mycota, volume 11. Springer, Berlin, 111–134 pp.
14. Butt T.M., Coates C.J., Dubovskiy I.M., Ratcliffe N.A., 2016. Entomopathogenic fungi: New insights into host-pathogen interactions. Advances in Genetics, 94, 307–364. https://doi.org/10.1016/bs.adgen.2016.01.006
15. Çakır C., 1992. Antalya ve çevresinde doğal olarak yetişen bazı bitkilerin fungitoksit potansiyellerinin araştırılması. M.S. thesis, Akdeniz University, 108 s., Antalya (in Turkish).
16. Çetin H., Erler F., Yanıkoğlu A., 2006. Toxicity of essential oils extracted from Origanum onites L. and Citrus aurentium L. against the pine processionary moth, Thaumetopoea wilkinsoni Tams. Folia biologica, 54 (3-4), 153-157. https://doi.org/10.3409/173491606778557473
17. Chakravarty I., Parmar V.M., Mandavgane, S.A. 2023. Current trends in essential oil (EO) production. Biomass Conversion and Biorefinery, 13, 15311–15334. https://doi.org/10.1007/s13399-021-01963-3
18. Eckard S., Bacher S., Enkerli J., Grabenweger G., 2017. A simple in vitro method to study interactions between soil insects, entomopathogenic fungi, and plant extracts. Entomologia Experimentalis et Applicata, 163, 315-327. https://doi.org/10.1111/eea.12578
19. Erdoğan P., Toros S., 2005. Melia azedarach L. (Meliaceae) ekstraktlarının Patates böceği [Leptinotarsa decemlineata Say (Col.: Chrysomelidae)] larvalarının gelişimi üzerine etkisi. Bitki Koruma Bülteni, 45 (1-4), 99-118.
20. Erler F., Ulug I., Yalcinkaya B., 2006. Repellent activity of five essential oils against Culex pipiens. Fitoterapia, 77, 491-494. https://doi.org/10.1016/j.fitote.2006.05.028
21. Gołębiowski M., Bojke A., Tkaczuk C., 2021. Effects of the entomopathogenic fungi Metarhizium robertsii, Metarhizium flavoviride, and Isaria fumosorosea on the lipid composition of Galleria mellonella larvae. Mycologia, 113 (3), 525–535. https://doi.org/10.1080/00275514.2021.1877520
22. Güdek M., Çetin H., 2017. Fumigant toxicity on adults of Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae) of essential oil from Rosmarinus officinalis L. and its side effects on chickpea grains. Journal of Essential Oil Bearing Plants, 20, 272-281. https://doi.org/10.1080/0972060X.2017.1281768
23. Hedin P.A., Hollingworth R.M., Masler E.P., Miyamoto J., Thompson D.G., 1997. Phytochemicals for pest control. ACS Symposium Series No 658, American Chemical Society, Washington DC, USA, 372 p.
24. Hongthong S., Wiparat N., Wongcharee S., Soythong P., 2025. Investigating the essential oil production from agricultural food waste: A case study Chaiyaphum province, Thailand. Case Studies in Chemical and Environmental Engineering, 12, 101286, https://doi.org/10.1016/j.cscee.2025.101286.
25. Hori M., 1998. Repellency of rosemary oil against Myzus persicae. in a laboratory and in a screenhouse. Journal of Chemical Ecology, 24, 1425–1432 https://doi.org/10.1023/A:1020947414051
26. Inglis G. D., Goettel M.S., Butt T. M., Strasser H., 2001. Use of hyphomycetous fungi for managing insect pests. In: Fungi as Biocontrol Agents: Progress, Problems and Potential. Butt T.M., Jackson C., Magan N. (Eds.). CAB International, 23-69 p.
27. Inglis G. D., Enkerli J., Goettel M.S., 2012. Chapter VII- Laboratory techniques used for entomopathogenic fungi: Hypocreales. In: Manual of Techniques in Invertebrate Pathology (Second Edition). Lacey L. A. (Ed.). Academic Press, 189-253 p. https://doi.org/10.1016/B978-0-12-386899-2.00007-5.
28. Isman M.B., 2000. Plant essential oils for pest and disease management. Crop Protection, 19 (8-10), 603–608. https://doi.org/10.1016/S0261-2194(00)00079-X
29. Isman M.B., 2006. Botanical insecticides, deterrents, and repellents, in modern agriculture and an increasingly regulated world. Annual Review of Entomology, 51, 45–6. https://doi.org/10.1146/annurev.ento.51.110104.151146
30. Isman M.B., Wilson J.A., Bradbury R., 2008. Insecticidal activities of commercial rosemary oils (Rosmarinus officinalis) against larvae of Pseudaletia unipuncta and Trichoplusia ni. in relation to their chemical composition. Pharmaceutical Biology, 46, 82–87. https://doi.org/10.1080/13880200701734661
31. Kalemba D., Kunicka A., 2003. Antibacterial and antifungal properties of essential oils. Current Medicinal Chemistry, 10 (10), 813–829. Doi: 10.2174/0929867033457719
32. Kanat M., Alma M.H., 2004. Insecticidal effects of essential oils from various plants against larvae of pine processionary moth (Thaumetopoea pityocampa Schiff) (Lepidoptera: Thaumetopoeidae). Pest Management Science, 60 (2), 173-177. DOI: 10.1002/ps.802
33. Keller S., Kessler P., Schweizer C., 2003. Distribution of insect pathogenic soil fungi in Switzerland with special reference to Beauveria brongniartii and Metarhizium anisopliae. Biocontrol, 48, 307–319. https://doi.org/10.1023/A:1023646207455
34. Kesdek M., Bayrak N., Kordali S., Usanmaz A., Contuk G., 2013. Larvicidal effects of some essenti al oils against larvae of the pine processionary moth (Thaumetopoea pityocampa [Denis & Schiffermüller]) (Lepidoptera: Thaumetopoeidae). Egyptian Journal of Biological Pest Control, 23, 201-207.
35. Kesdek M., Kordali S., Çoban K., Usanmaz A., Ercişli S., 2014. Larvicidal effect of some plant extracts on the pine processionary moth, Thaumetopoea pityocampa (Denis & Schiffermuller) in laboratory conditions. Acta Scientiarum Polonorum Hortorum Cultus, 13 (5), 145-162.
36. Kesdek M., Kordali S., Usanmaz A., Ercişli S., 2015. The toxicity of essential oils of some plant species against adults of Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae). Comptes Rendes de l ̍Académie Bulgare des Sciences, 68 (1), 127-136.
37. Khalfi O., Sahraoui N., Bentahar F., Boutekedjiret C., 2008. Chemical composition and insecticidal properties of Origanum glandulosum (Desf.) essential oil from Algeria. Journal of the Science of Food and Agriculture, 88, 1562–1566. https://doi.org/10.1002/jsfa.3251
38. Klingen I., Eilenberg J., Meadow R., 2002. Effects of farming system, field margins and bait insect on the occurrence of insect pathogenic fungi in soils. Agriculture Ecosystems and Environment, 91 (1-3), 191–198. https://doi.org/10.1016/S0167-8809(01)00227-4
39. Koul O., WalIa S., Dhaliwal G.S., 2008. Essential Oils as Green Pesticides: Potential and Constraints. Biopesticides International, 4 (1), 63–84.
40. Kordali S., Aslan İ., Çalmaşur Ö., Çakır A., 2006. Toxicity of essential oils isolated from three Artemisia species and some of their major components to granary weevil, Sitophilus granarius (L.) (Coleoptera: Curculionidae). Industrial Crops and Products, 23 (2), 162-170. https://doi.org/10.1016/j.indcrop.2005.05.005
41. Kordali S., Çakır A., Özer H., Çakmakçı R., Kesdek M., 2008. Antifungal, phytotoxic and insecticidal properties of essential oil isolated from Turkish Origanum acutidens and its three components, carvacrol, thymol and p-cymene. Bioresource Technology, 99 (18), 8788-8795. https://doi.org/10.1016/j.biortech.2008.04.048
42. Koschier E. H., Sedy K. A., 2003. Labiate essential oils affecting host selection and acceptance of Thrips tabaci. Lindeman. Crop Protection, 22, 929–934. https://doi.org/10.1016/S0261-2194(03)00092-9
43. Kumar S., Singh A., Kumar M., 2022. Biopesticides: An alternative to synthetic pesticides. Environmental Sustainability, 5 (3), 263–274. https://doi.org/10.1007/s42398-022-00236-8
44. Lacey L. A., Frutos R., Kaya H. K., Vail P., 2001. Insect pathogens as biological control agents: Do they have a future? Biological Control, 21 (3), 230-248. https://doi.org/10.1006/bcon.2001.0938
45. Lak F., Zandi-Sohani N., Ghodoum Parizipour M.H., Ebadollahi A., 2022. Synergic effects of some plant-derived essential oils and Iranian isolates of entomopathogenic fungus Metarhizium anisopliae Sorokin to control Acanthoscelides obtectus (Say) (Coleoptera: Chrysomelidae). Frontiers in Plant Science, 13, 1075761. https://doi.org/10.3389/fpls.2022.1075761
46. Lucia A, Audino P.G., Seccacini E., Licastro S., Zerba E., Masuh H., 2007. Larvicidal effect of Eucalyptus grandis essential oil and turpentine and their major components on Aedes aegypti larvae. Journal of the American Mosquito Control Association, 23, 299-303. https://doi.org/10.2987/8756-971X(2007)23[299:LEOEGE]2.0.CO;2
47. Meyling N. V., Eilenberg J., 2007. Ecology of the entomopathogenic fungi Beauveria bassiana and Metarhizium anisopliae in temperate agroecosystems: Potential for conservation biological control. Biological Control, 43 (1), 145–155. https://doi.org/10.1016/j.biocontrol.2007.07.007
48. Mudrončeková S., Ferenčík J., Gruľová D., Barta M., 2019. Insecticidal and repellent effects of plant essential oils against Ips typographus. Journal of Pest Science, 92, 595–608. https://doi.org/10.1007/s10340-018-1038-1
49. Ortiz-Urquiza, A., Keyhani N. O., 2013. Action on the surface: Entomopathogenic fungi versus the insect cuticle. Insects, 4 (3), 357–374. https://doi.org/10.3390/insects4030357
50. Papachristos D.P., Karamanoli K.I., Stamopoulos D.C, Menkissoglu-Spiroudi U., 2004. The relationship between the chemical composition of three essential oils and their insecticidal activity against Acanthoscelides obtectus. (Say). Pest Management Science, 60, 514–520. https://doi.org/10.1002/ps.798
51. Rajendran S., Sriranjini V., 2008. Plant products as fumigants for stored-product insect control. Journal of Stored Products Research, 44, 126-135. https://doi.org/10.1016/j.jspr.2007.08.003
52. Razmjou J., Davari M., Ebadollahi A., 2016. Effect of two plant essential oils and the entomopathogenic fungus, Lecanicillium muscarium (zare & gams) on the cotton aphid, Aphis gossypii Glover. Egyptian Journal of Biological Pest Control, 26 (4), 775-779.
53. Regnault-Roger C., Hamraoui A., Holeman M., Theron E., Pinel R., 1993. Insecticidal effect of essential oils from Mediterranean plant upon Acanthoscelides obtectus Say (Coleoptera: Bruchidae), a pest of kidney bean (Phaseolus vulgaris). Journal of Chemical Ecology, 19, 1233-1244.
54. Rocha I.U., Bitencourt R.O.B., Freitas A.M., Moreira H.V.S., Magalhães K.L.A., de Souza B.A., Golo P.S., Chaves D.S.A, Bittencourt V.R.E.P., Angelo I.D., 2024. Exploiting the combination of entomopathogenic fungi and Illicium verum essential oil against Aedes aegypti larvae. Biological Control, 193, 105526, https://doi.org/10.1016/j.biocontrol.2024.105526.
55. Rozman V., Kalinovic I., Korunic Z., 2007. Toxicity of naturally occurring compounds of Lamiaceae and Lauraceae to three stored-product insects. Journal of Stored Products Research, 43, 349–355. https://doi.org/10.1016/j.jspr.2006.09.001
56. Rui C., Abelho A., Ribeiro N., 2018. A Review of the scientific literature for optimal conditions for mass rearing Tenebrio molitor (Coleoptera: Tenebrionidae). Journal of Entomological Science, 53 (4), 434-454. DOI: 10.18474/JES17-67.1
57. Shaaya E., Ravyd U., Paster N., Kostjukovsky M., Menasherov M., 1993. Essential oils and their components as active fumigants against several species of stored product insects and fungi. Journal of Chemical Ecology, 344, 131-137.
58. Shah P.A., Pell J.K., 2003. Entomopathogenic fungi as biological control agents. Applied Microbiology and Biotechnology, 61 (5–6), 413–423. https://doi.org/10.1007/s00253-003-1240-8
59. Soković M., van Griensven L.J., 2006. Antimicrobial activity of essential oils and their components against the three major pathogens of the cultivated button mushroom, Agaricus bisporus. European Journal of Plant Pathology 116, 211–224. https://doi.org/10.1007/s10658-006-9053-0
60. Soylu E.M., Soylu S., Kurt Ş., 2006. Antimicrobial activity of essential oils the essential oils of various plants against tomato late blight disease agent Phytophthora infestans. Mycopathologia, 161 (2), 119-28. doi: 10.1007/s11046-005-0206-z.
61. Soylu S, Yigitbas H, Kurt Ş., 2007. Antifungal effects of essential oils from oregano and fennel on Sclerotinia sclerotiorum. Journal of Applied Microbiology, 103 (4), 1021–1030. DOI: 10.1111/j.1365-2672.2007.03310.x
62. Tinkeu N.L.S., Goudoum A, Ngassoum M.B., Mapongmetsem P.M., Kouninki H., Hance T., 2004. Persistence of the insecticidal activity of five essential oils on the maize weevil Sitophilus zeamais (Motsch.) (Coleoptera: Curculionidae). Communications in Agricultural and Applied Biological Sciences, 69 (3), 145-147.
63. Toshova T., Velchev D., Barta M., 2022. Insecticide activity of Greek oregano essential oil and entomopathogenic fungus Metarhizium pemphigi against Diabrotica virgifera virgifera LeConte. Cereal Research Communications, 50, 1045–1054. https://doi.org/10.1007/s42976-022-00266-1
64. Tunç I., Berger B.M., Erler F., Dagli F., 2000. Ovicidal activity of essential oils from five plants against two stored-products insects. Journal of Stored Products Research, 36 (2), 161–161. DOI: 10.1016/S0022-474X(99)00036-3
65. Türkölmez S., Soylu E.M., 2014. Antifungal efficacies of plant essential oils and main constituents against soil-borne fungal disease agents of bean. Journal of Essential Oil Bearing Plants, 17(2), 203–211. DOI: 10.1080/0972060X.2014.895160
66. Usanmaz Bozhüyük A., Kesdek M., Kordali Ş., Özcan S., 2018. Artemisia santanicum L. ve Artemisia absinthium L. uçucu yağlarının Çam Keseböceği (Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera: Thaumetopoeidae)’ne larvasidal etkisi. Journal of the Institute of Science and Technology, 8(3), 63-70. https://doi.org/10.21597/jist.458575
67. Usanmaz Bozhüyük A., Kordali Ş., 2019. Ocimum Basilicum L. uçucu yağının antifungal etkileri. Erciyes Tarım ve Hayvan Bilimleri Dergisi, 2(1), 45-57. https://izlik.org/JA46TP86ZB
68. Üstüner T., Kordali Ş., Usanmaz Bozhüyük A., Kesdek M., 2018. Investigation of pesticidal activities of essential oil of Eucalyptus camuldulensis Dehnh. Records of Natural Products 12 (6), 557-568. https://doi.org/10.25135/rnp.64.18.02.088
69. van Lenteren J.C., Bolckmans K., Köhl J., Ravensberg W. J., Urbaneja A., 2017. Biological control using invertebrates and microorganisms: Plenty of new opportunities. BioControl, 63 (1), 39–59. https://doi.org/10.1007/s10526-017-9801-4
70. Wu J, Yang B, Zhang X, Cuthbertson AGS, Ali S., 2021. Synergistic Interaction between the Entomopathogenic Fungus Akanthomyces attenuatus (Zare & Gams) and the Botanical Insecticide Matrine against Megalurothrips usitatus (Bagrall). Journal of Fungi, 7 (7), 536. https://doi.org/10.3390/jof7070536
71. Yıldırım E., Kesdek M., Kordali S., 2005. Effects of essential oil of three plant species on Tribolium confusum du Val and Sitophilus granarius (L.) (Coleoptera: Tenebrionidae and Curculionidae). Fresenius Environmental Bulletin, 14, 574-578.
72. Zimmermann G., 2007a. Review on safety of the entomopathogenic fungus Metarhizium anisopliae. Biocontrol Science and Technology, 17 (9), 879–920. https://doi.org/10.1080/09583150701593963
73. Zimmermann G., 2007b. Review on safety of the entomopathogenic fungi Beauveria bassiana and Beauveria brongniartii. Biocontrol Science and Technology, 17 (6), 553–596. https://doi.org/10.1080/09583150701309006