First report of the destroyer ant Trichomyrmex destructor (Jerdon, 1851) (Hymenoptera, Formicidae, Myrmicinae) as a pest of red dragon fruit Selenicereus costaricensis (Cactaceae): a case of extrafloral nectaries, evolved for self-defense, ended up becoming a source of harm to the plant

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First report of the destroyer ant Trichomyrmex destructor (Jerdon, 1851) (Hymenoptera, Formicidae, Myrmicinae) as a pest of red dragon fruit Selenicereus costaricensis (Cactaceae): a case of extrafloral nectaries, evolved for self-defense, ended up becoming a source of harm to the plant | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Short Report First report of the destroyer ant Trichomyrmex destructor (Jerdon, 1851) (Hymenoptera, Formicidae, Myrmicinae) as a pest of red dragon fruit Selenicereus costaricensis (Cactaceae): a case of extrafloral nectaries, evolved for self-defense, ended up becoming a source of harm to the plant P. Akheela, K. D. Prathapan, T. Santhoshkumar This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4733344/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract The destroyer ant, Trichomyrmex destructor (Jerdon, 1805) (Formicidae, Myrmicinae), is newly reported as a pest of the red dragon fruit Selenicereus costaricensis (F. A. C. Weber) S. Arias & N. Korotkova ex Hammel (Cactaceae). It feeds on the areoles, scale leaves and surrounding green tissues at the base of spines of S. costaricensis , due to the presence of extrafloral nectaries. The damaged portions turned yellow and then dark brown to black. The extent of infestation on the cladode was 0–50%. Besides cladode, flowers and fruits were also damaged. Up to 90% fruits and/flowers were infested on four clumps. This is a case of direct cost of production of extrafloral nectaries, which are protective in function against herbivores. This is the first report of extrafloral nectaries in S. costaracensis. Selenicereus costaricensis India Cactaceae herbivory Figures Figure 1 Figure 2 Figure 3 INTRODUCTION The destroyer ant, Trichomyrmex destructor (Jerdon) (Myrmicinae) (Figs. 1–3), originally described from India by Jerdon in 1851 in the genus Atta Fabricius, 1805 and probably of north African origin (Wetterer 2009 ), is a tramp species dispersed through trade. It is widely distributed in the tropics and subtropics of the Old World; Florida in the United States, the West Indies and the Galapagos Islands in the New World; and Australia. Wetterer ( 2009 ) has listed 107 countries and islands in its range of distribution. Bolton ( 1987 ) provided description of T. destructor and Wetterer ( 2009 ) gave salient identification characters. Workers of T. destructor are polymorphic, varying greatly in size displaying allometric variation and markedly bicolored – light yellow to yellow-brown, except the gaster (Fig. 3) dark brown to nearly black in distal two-third. They can be readily identified by the finely transversely striate vertex and dorsum of propodeum as well as the deep metanotal groove (Fig. 2). Antenna is 12 segmented, including a three segmented club with increasing size towards apex (Fig. 1). Eyes small with 4–6 ommatidia in longest row. Mandibles are 4-dentate and paired longitudinal carinae on clypeus obscure (Fig. 1). Petiole higher and less broadly rounded and narrower than postpetiole (Fig. 2). Dorsum with erect setae. Jerdon ( 1851 ), in his original description of T. destructor , also recorded its natural history: “ They live in holes in the ground, or in walls and are very numerous in individuals. They prefer animal to vegetable substance, destroying dead insects, bird skins etc., but also feed greedily on sugar. They are common in all parts of India, and often prove very troublesome and destructive to the naturalist. ” Trichomyrmex destructor is a generalist with a broad diet of living and dead insects, carbohydrates, nectar and seeds (Bolton 1987 ; Harris et al., 2005 ). Trichomyrmex destructor is a major urban pest that damages property, attacking people, kill caged animals and carry human pathogens (Harris et al., 2005 ). They also forage and nest in and around agricultural crops and other vegetation. They forage on the crown, trunk as well as base of coconut trees (Way et al., 1989 ) and nest in trees in citrus orchards (Collingwood et al., 1997 ), base of Acacia sp., near the date palm Phoenix dactylifera L. and Calotropis procera W. T. Aiton (Sharaf et al., 2016 ). Kavya ( 2023 ) has reported direct feeding damage by T. destructor on okra ( Abelmoschus esculentus (L.) Moench; Malvaceae) and eggplant ( Solanum melongena L.; Solanaceae). Here we report direct damage by T. destructor on the red dragon fruit Selenicereus costaricensis (F. A. C. Weber) S. Arias & N. Korotkova ex Hammel (Cactaceae). Dragon fruit is a group of vine cacti belonging to the genera Selenicereus Britton & Rose and Hylocereus Britton & Rose, native to the Central and South America. These are perennial, clambering, fast growing members of Cactaceae having 3–5 ridged stems (Fig. 4), widely cultivated in the tropics and subtropics for its fruit. Introduced to India in the late 1990s’, dragon fruit is cultivated in 3000–4000 ha across the country, with an annual production of about 12,000 tons of fruit in 2020 (Wakchure et al., 2021). Being tolerant to stress and adapted to arid and barren lands, a tenfold increase in its cultivation in India by 2025 is predicted (Wakchure et al., 2021). Dragon fruit in India is generally free of insect pests, except undetermined species of ants, scale insects and mealy bugs (Wakchure et al., 2021), tobacco cutworm, Spodoptera litura (Fabricius) (Prathapan & Santhoshkumar, 2022 ) and the fruit flies B. correcta (Bezzi) (Wakchure et al., 2021) and Bactrocera dorsalis (Hendel) (Wakchure et al., 2021; Akheela et al., 2024 ). MATERIAL AND METHODS Infestation of T. destructor on the red dragon fruit or red pitahaya S. costaricensis was observed at five localities in Thiruvananthapuram and Kollam Districts in Kerala during 2022–2024 (Table 1 ). The nature and extent of damage were recorded. Images of the ant in the laboratory were captured using a Canon EOS 700 DSLR camera mounted on a Leitz compound microscope. Custom-made illuminators and diffusers were used for lighting. Photographs in the field were taken with a Laowa 2x lens mounted on a Nikon D 3000 SLR camera. Simpex 666 Speed Light with a custom-made diffuser was used for lighting. Voucher specimens of T. destructor are deposited in the National Bureau of Agricultural Insect Resources, Bengaluru (Accession number NIM/NBAIR/HYM/FORM/24624). Table 1 Infestation of Trichomyrmex destructor on Selenicereus costaricensis Location GPS coordinates Date of observation Stage of crop No of infested clumps Damage Fruit/flower Cladode (mean) Thiruvananthapuram District Pangode Latitude 8.7744009 N, Longitude 76.9928065 E April 2022 Vegetative 6 0 30/119 (25%) Vembayam Latitude 8.6318480 N, Longitude 76.9533113 E September 2023 Flowering and fruiting 4 Observed 16/207 (8%) Maranallur Latitude 8.4679820 N, Longitude 76.0424700 E 10 June 2024 Flowering & fruiting 4 45/50 (90%) 0 Kollam District Karingannur Latitude 8.429738 N, Longitude 76.985509 E 30 April 2024 Flowering & fruiting 4 Observed 0 Kundara Latitude 8.9785760 N, Longitude 76.6776440 E 30 April 2024 Vegetative 1 0 6/12 (50%) RESULTS At two locations, the crop was in vegetative stage, while the plants were at the bearing stage at the other three (Table 1 ). At Pangode, six clumps, each having 17–46 cladodes were infested. Of the 119 cladodes observed, 30 were infested. The extent of infestation varied from 21.2–35.2% (mean 25.2%). At Vembayam, four clumps, each having 33–67 cladodes, were infested. Of the 207 cladodes observed, 16 were damaged. The extent of infestation varied from 4.6–11.9% (mean 7.7%). At Kundara, six cladodes out of twelve (50%) in a single clump, in a homestead, were observed affected. The infestation was also observed at Karingannur in Kollam on flowers and fruits, however, cladodes were free of ant damage. Besides cladode, flowers and fruits were also affected at Vembayam. At Maranallur, 45 out of 50 (90%) fruits and/ flowers on four infested clumps were damaged, however, no damage was observed on the cladodes. Ants ate the areole (Figs. 4, 5) and the surrounding tissues at the base of spines, resulting in characteristic damage (Figs. 6, 7). Due to feeding of the tissues at the base of areole, deep emarginations were formed on the ridges of the cladode, in the place of spines (Fig. 6). The damaged portions turned yellow and then dark brown to black (Figs. 6, 7). The affected plants appeared weak and their growth was suppressed. At Vembayam, besides the base of spines, they also fed on flowers (Figs. 8, 9) and developing green fruits (Fig. 10) as well as mature, ripening fruits (Fig. 11). Ants fed directly on the flowers (Fig. 9) and fruits (Figs. 10, 11), resulting in sunken areas. On fruits, these sunken areas turned black (Figs. 10, 11). Infested fruits became visually unattractive and lost their market value. DISCUSSION In Cactaceae, 156 species in 40 genera are known to have extrafloral nectaries (Keeler et al., 2015–2024). Selenicereus undatus (Haw.) D. R. Hunt, the white-fleshed pitahaya, possesses extrafloral nectaries (Keeler et al., 2015–2024). It was observed that S . costaricensis also have nectarines within the areole at the base of the spines as well as under the small, scale-like leaves beneath the areole on developing cladodes (Fig. 5). It is the extrafloral nectaries that attract T. destructor to the dragon fruit cactus, which, according to Jerdon ( 1851 ), ‘ feeds greedily on sugar’. Extrafloral nectaries attract aggressive arthropods that protect developing leaves, shoots and flowers from herbivores and thus form an important weapon in the vast repertoire of plant defense mechanisms (Marazzi et al., 2013 ; Heil, 2015 ). However, greater defense is accompanied by indirect costs such as reduced competitive ability, deterrence of pollinators (Ness, 2006 ) and natural enemies of herbivores (Callejas-Chavero et al. 2020 ) or exploitation of extrafloral nectaries by herbivores to their advantage when they are attracted to and retained by the extrafloral nectaries (Turlings & Wӓckers, 2004 ; Popa, 2020). Selective, targeted feeding on extrafloral nectaries by several insect species, including some that are otherwise not primarily herbivorous, are known (Gish et al., 2015 ). This is an extreme case of direct cost of production of extrafloral nectaries, which are supposed to be protective in function against herbivores. Here the ‘bodyguards’ of the plant turn assailants, by shifting from predation to herbivory. Flowers and fruits, when present, were preferred by the ants to the cladodes. This is probably due to the availability of more sugars and carbohydrates in fruits and flowers. This is the first report of T. destructor as a pest of S. costaracensis and the first report of extrafloral nectaries in S. costaracensis . Declarations Author Contribution Author contributions: AP independently collected data and made observations; ST conducted fieldwork, collected data and recorded observations; KDP conceptualized the study, carried out fieldwork, made observations and prepared the first draft of the manuscript. All authors reviewed and revised the manuscript. Acknowledgement We are indebted to Sahana Shree, Ashoka Trust for Research in Ecology and the Environment (ATREE), Bengaluru for the identification of T. destructor. Sandra Maria Mathew prepared Figs 1 – 3. This study was partially funded by the Kerala Agricultural University. Anooj S. S. critically reviewed the manuscript. References Akheela, P., Santhoshkumar, T., & Varghese, T. S. (2024). First report of Bactrocera dorsalis (Hendel) (Diptera, Tephritidae) on the white-fleshed dragon fruit Selenicereus undatus (Haworth) DR Hunt (Cactaceae) in India. Entomon , 49(2), 283-286. https://doi.org/10.33307/entomon.v49i2.1190 Bolton, B. (1987). A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr (Hymenoptera: Formicidae). Bulletin of the British Museum (Natural History). Entomology 54(3), 263-452. https://biostor.org/reference/113868 Callejas-Chavero, A., Martínez-Hernández, D., Flores-Martínez, A., Moncada-Orellana, A., Diaz-Quiñones, Y., & Vargas-Mendoza, C. F. (2020). Herbivory in Cacti: Fitness Effects of Two Herbivores, One Tending Ant on Myrtillocactus geometrizans (Cactaceae). In: Núñez-Farfán, J., Valverde, P. (eds) Evolutionary Ecology of Plant-Herbivore Interaction. (pp. 109-134). Springer, Cham . Collingwood, C. A., Tigar, B. J., & Agosti, D. (1997). Introduced ants in the United Arab Emirates. Journal of Arid Environments 37(3), 505-512. https://doi.org/10.1006/jare.1997.0309 Gish, M., Mescher, M. C., & DeMoraes, C. M. (2015). Targeted predation of extrafloral nectaries by insects despite localized chemical defences. Proceedings of the Royal Society B: Biological Sciences 282(1816): 20151835. https://doi.org/10.1098/rspb.2015.1835 Harris, R., Abbott, K., Barton, K., Berry, J., Don, W., Gunawardana, D., Lester, P., Rees, J., Stanley M., Sutherland, A., & Toft, R. (2005). Invasive ant pest risk assessment project for Biosecurity New Zealand. Series of unpublished Landcare Research contract reports to Biosecurity New Zealand. BAH/35/2004-1. Heil, M. (2015). Extrafloral nectar at the plant-insect interface: a spotlight on chemical ecology, phenotypic plasticity, and food webs. Annual Review of Entomology 60: 213-232. https://doi.org/10.1146/annurev-ento-010814-020753 Jerdon, T. C. (1851). A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science 17, 103-127. Kavya, K. T. (2023). Management of phytophagous ants in vegetable crops under homestead cultivation. (Master’sThesis). Kerala Agricultural University, Vellayani. 83 p. Keeler, K. H., Porturas, L. D., & Weber, M. G. (2015–2024). World list of plants with extrafloral nectaries. www.extrafloralnectaries.org. Accessed on 18 April 2024 Marazzi, B., Bronstein, J. L., & Koptur, S. (2013). The diversity, ecology and evolution of extrafloral nectaries: current perspectives and future challenges. Annals of Botany 111(6),1243-1250. https://doi.org/10.1093/aob/mct109 Ness, J. H. (2006). A mutualism's indirect costs: the most aggressive plant bodyguards also deter pollinators. Oikos 113(3), 506-514. https://doi.org/10.1111/j.2006.0030-1299.14143.x Popa, V. I., Georgescu, I.M., Potor, C. D., & Hoza, D. (2020). Defensive role of extrafloral nectaries. Scientific Papers. Series B. Horticulture 64(1), 693-700. Prathapan, K. D., & Santhoshkumar, T. (2022). First report of Spodoptera litura (Fabricius) (Noctuidae) on the dragon fruit species Selenicereus costaricensis (F.A.C.Weber) S.Arias & N.Korotkova and Selenicereus megalanthus (K.Schum. ex Vaupel) Moran (Cactaceae). Journal of the Lepidopterists’ Society 76 (4), 282-283. https://doi.org/10.18473/lepi.76i4.a9 Sharaf, M. R., Salman, S., Al Dhafer, H. M., Akbar, S. A., Abdel-Dayem, M. S., & Aldawood, A. S. (2016). Taxonomy and distribution of the genus Trichomyrmex Mayr, 1865 (Hymenoptera: Formicidae) in the Arabian Peninsula, with the description of two new species . European Journal of Taxonomy 246,1-36. https://doi.org/10.5852/ejt.2016.246 Turlings, T. C. J., &Wӓckers, F. (2004). Recruitment of predators and parasitoids by herbivore-injured plants. In: Advances in Insect Chemical Ecology (R. T. Cardé, ed.). (pp. 21-75). Cambridge University Press. Wakchaure, G. C., Satish Kumar, Meena, K. K., Rane, J., & Pathak, H. (2021). Dragon Fruit Cultivation in India: Scope, Constraints and Policy Issues. Technical Bulletin No 27,47. ICAR–National Institute of Abiotic Stress Management, Baramati, Pune, Maharashtra, India. Way, M. J., Cammell, M. E., Bolton, B., & Kanagaratnam, P. (1989). Ants (Hymenoptera: Formicidae) as egg predators of coconut pests, especially in relation to biological control of the coconut caterpillar, Opisina arenosella Walker (Lepidoptera: Xyloryctidae), in Sri Lanka. Bulletin of Entomological Research 79(2), 219-234. https://doi.org/10.1017/S0007485300018204 Wetterer, J. K. (2009). Worldwide spread of the destroyer ant, Monomorium destructor (Hymenoptera: Formicidae). Myrmecological News 12, 97-108. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-4733344","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Short Report","associatedPublications":[],"authors":[{"id":332274244,"identity":"c6153108-cbb9-4fcf-ba86-f318b49993f4","order_by":0,"name":"P. 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Santhoshkumar","email":"","orcid":"","institution":"Kerala Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"T.","middleName":"","lastName":"Santhoshkumar","suffix":""}],"badges":[],"createdAt":"2024-07-13 04:23:16","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4733344/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4733344/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":62324416,"identity":"6d87d118-b467-4a3d-89e5-7bbbd80c1601","added_by":"auto","created_at":"2024-08-13 02:30:04","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4427719,"visible":true,"origin":"","legend":"\u003cp\u003eFigs 1 – 3. \u003cem\u003eTrichomyrmex destructor \u003c/em\u003e(scale bars = 0.2 mm)\u003cem\u003e. \u003c/em\u003e1. head, 2. mesosoma, petiole and postpetiole in dorsal view, 3. gaster.\u003c/p\u003e","description":"","filename":"PLATE1.png","url":"https://assets-eu.researchsquare.com/files/rs-4733344/v1/29b7cd366b62aaf14fde4511.png"},{"id":62324415,"identity":"e25aaf82-0d8c-4791-a693-2b5ca8e39707","added_by":"auto","created_at":"2024-08-13 02:30:04","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":10977368,"visible":true,"origin":"","legend":"\u003cp\u003eFigs 4 – 7. \u003cem\u003eSelenicereus costaricensis. \u003c/em\u003e4. developing cladode, 5. areoles, note scale leaf beneath (indicated by arrow), 6, 7. areole and surrounding tissues eaten away by ants.\u003c/p\u003e","description":"","filename":"Plate3.png","url":"https://assets-eu.researchsquare.com/files/rs-4733344/v1/42b58af055037cecbdf2a734.png"},{"id":62324414,"identity":"3a40edb9-4bcd-45be-b33e-bb4916eed837","added_by":"auto","created_at":"2024-08-13 02:30:04","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":10255885,"visible":true,"origin":"","legend":"\u003cp\u003eFigs 8 – 11. \u003cem\u003eSelenicereus costaricensis. \u003c/em\u003e8. flower bud infested by ants, 9. closeup of damage on flower bud, 10. infestation on developing fruit, note the black color on the rind damaged by ants, 11. infestation on ripening fruit, note the black color on the damaged surface.\u003c/p\u003e","description":"","filename":"Plate2.png","url":"https://assets-eu.researchsquare.com/files/rs-4733344/v1/e2e8814f59df5bf4fbc05ebd.png"},{"id":62775454,"identity":"d455ed8c-cb99-4295-adb3-20eeb7ea06f4","added_by":"auto","created_at":"2024-08-19 10:15:32","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":43952288,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4733344/v1/8ae1a69e-2925-4e8f-99a4-aee98c657001.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"First report of the destroyer ant Trichomyrmex destructor (Jerdon, 1851) (Hymenoptera, Formicidae, Myrmicinae) as a pest of red dragon fruit Selenicereus costaricensis (Cactaceae): a case of extrafloral nectaries, evolved for self-defense, ended up becoming a source of harm to the plant","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eThe destroyer ant, \u003cem\u003eTrichomyrmex destructor\u003c/em\u003e (Jerdon) (Myrmicinae) (Figs.\u0026nbsp;1\u0026ndash;3), originally described from India by Jerdon in 1851 in the genus \u003cem\u003eAtta\u003c/em\u003e Fabricius, 1805 and probably of north African origin (Wetterer \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), is a tramp species dispersed through trade. It is widely distributed in the tropics and subtropics of the Old World; Florida in the United States, the West Indies and the Galapagos Islands in the New World; and Australia. Wetterer (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) has listed 107 countries and islands in its range of distribution.\u003c/p\u003e \u003cp\u003eBolton (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1987\u003c/span\u003e) provided description of \u003cem\u003eT. destructor\u003c/em\u003e and Wetterer (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) gave salient identification characters. Workers of \u003cem\u003eT. destructor\u003c/em\u003e are polymorphic, varying greatly in size displaying allometric variation and markedly bicolored \u0026ndash; light yellow to yellow-brown, except the gaster (Fig.\u0026nbsp;3) dark brown to nearly black in distal two-third. They can be readily identified by the finely transversely striate vertex and dorsum of propodeum as well as the deep metanotal groove (Fig.\u0026nbsp;2). Antenna is 12 segmented, including a three segmented club with increasing size towards apex (Fig.\u0026nbsp;1). Eyes small with 4\u0026ndash;6 ommatidia in longest row. Mandibles are 4-dentate and paired longitudinal carinae on clypeus obscure (Fig.\u0026nbsp;1). Petiole higher and less broadly rounded and narrower than postpetiole (Fig.\u0026nbsp;2). Dorsum with erect setae.\u003c/p\u003e \u003cp\u003eJerdon (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1851\u003c/span\u003e), in his original description of \u003cem\u003eT. destructor\u003c/em\u003e, also recorded its natural history: \u0026ldquo;\u003cem\u003eThey live in holes in the ground, or in walls and are very numerous in individuals. They prefer animal to vegetable substance, destroying dead insects, bird skins etc., but also feed greedily on sugar. They are common in all parts of India, and often prove very troublesome and destructive to the naturalist.\u003c/em\u003e\u0026rdquo; \u003cem\u003eTrichomyrmex destructor\u003c/em\u003e is a generalist with a broad diet of living and dead insects, carbohydrates, nectar and seeds (Bolton \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Harris et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eTrichomyrmex destructor\u003c/em\u003e is a major urban pest that damages property, attacking people, kill caged animals and carry human pathogens (Harris et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). They also forage and nest in and around agricultural crops and other vegetation. They forage on the crown, trunk as well as base of coconut trees (Way et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1989\u003c/span\u003e) and nest in trees in citrus orchards (Collingwood et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), base of \u003cem\u003eAcacia\u003c/em\u003e sp., near the date palm \u003cem\u003ePhoenix dactylifera\u003c/em\u003e L. and \u003cem\u003eCalotropis procera\u003c/em\u003e W. T. Aiton (Sharaf et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Kavya (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) has reported direct feeding damage by \u003cem\u003eT. destructor\u003c/em\u003e on okra (\u003cem\u003eAbelmoschus esculentus\u003c/em\u003e (L.) Moench; Malvaceae) and eggplant (\u003cem\u003eSolanum melongena\u003c/em\u003e L.; Solanaceae). Here we report direct damage by \u003cem\u003eT. destructor\u003c/em\u003e on the red dragon fruit \u003cem\u003eSelenicereus costaricensis\u003c/em\u003e (F. A. C. Weber) S. Arias \u0026amp; N. Korotkova ex Hammel (Cactaceae).\u003c/p\u003e \u003cp\u003eDragon fruit is a group of vine cacti belonging to the genera \u003cem\u003eSelenicereus\u003c/em\u003e Britton \u0026amp; Rose and \u003cem\u003eHylocereus\u003c/em\u003e Britton \u0026amp; Rose, native to the Central and South America. These are perennial, clambering, fast growing members of Cactaceae having 3\u0026ndash;5 ridged stems (Fig.\u0026nbsp;4), widely cultivated in the tropics and subtropics for its fruit. Introduced to India in the late 1990s\u0026rsquo;, dragon fruit is cultivated in 3000\u0026ndash;4000 ha across the country, with an annual production of about 12,000 tons of fruit in 2020 (Wakchure et al., 2021). Being tolerant to stress and adapted to arid and barren lands, a tenfold increase in its cultivation in India by 2025 is predicted (Wakchure et al., 2021). Dragon fruit in India is generally free of insect pests, except undetermined species of ants, scale insects and mealy bugs (Wakchure et al., 2021), tobacco cutworm, \u003cem\u003eSpodoptera litura\u003c/em\u003e (Fabricius) (Prathapan \u0026amp; Santhoshkumar, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and the fruit flies \u003cem\u003eB. correcta\u003c/em\u003e (Bezzi) (Wakchure et al., 2021) and \u003cem\u003eBactrocera dorsalis\u003c/em\u003e (Hendel) (Wakchure et al., 2021; Akheela et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e"},{"header":"MATERIAL AND METHODS","content":"\u003cp\u003eInfestation of \u003cem\u003eT. destructor\u003c/em\u003e on the red dragon fruit or red pitahaya \u003cem\u003eS. costaricensis\u003c/em\u003e was observed at five localities in Thiruvananthapuram and Kollam Districts in Kerala during 2022\u0026ndash;2024 (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The nature and extent of damage were recorded. Images of the ant in the laboratory were captured using a Canon EOS 700 DSLR camera mounted on a Leitz compound microscope. Custom-made illuminators and diffusers were used for lighting. Photographs in the field were taken with a Laowa 2x lens mounted on a Nikon D 3000 SLR camera. Simpex 666 Speed Light with a custom-made diffuser was used for lighting. Voucher specimens of \u003cem\u003eT. destructor\u003c/em\u003e are deposited in the National Bureau of Agricultural Insect Resources, Bengaluru (Accession number NIM/NBAIR/HYM/FORM/24624).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eInfestation of \u003cem\u003eTrichomyrmex destructor\u003c/em\u003e on \u003cem\u003eSelenicereus costaricensis\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c2\" namest=\"c1\" rowspan=\"2\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eGPS coordinates\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eDate of observation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eStage of crop\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eNo of infested clumps\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c8\" namest=\"c7\"\u003e \u003cp\u003eDamage\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFruit/flower\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eCladode\u003c/p\u003e \u003cp\u003e(mean)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eThiruvananthapuram District\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePangode\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLatitude 8.7744009 N, Longitude 76.9928065 E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eApril 2022\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVegetative\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e30/119\u003c/p\u003e \u003cp\u003e(25%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVembayam\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLatitude 8.6318480 N, Longitude 76.9533113 E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSeptember 2023\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFlowering and fruiting\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eObserved\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e16/207\u003c/p\u003e \u003cp\u003e(8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMaranallur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLatitude 8.4679820 N, Longitude 76.0424700 E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10 June 2024\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFlowering \u0026amp; fruiting\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e45/50\u003c/p\u003e \u003cp\u003e(90%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eKollam District\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eKaringannur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLatitude 8.429738 N, Longitude 76.985509 E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30 April 2024\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFlowering \u0026amp; fruiting\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eObserved\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eKundara\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLatitude 8.9785760 N, Longitude 76.6776440 E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30 April 2024\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVegetative\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6/12\u003c/p\u003e \u003cp\u003e(50%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"RESULTS","content":"\u003cp\u003eAt two locations, the crop was in vegetative stage, while the plants were at the bearing stage at the other three (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). At Pangode, six clumps, each having 17\u0026ndash;46 cladodes were infested. Of the 119 cladodes observed, 30 were infested. The extent of infestation varied from 21.2\u0026ndash;35.2% (mean 25.2%). At Vembayam, four clumps, each having 33\u0026ndash;67 cladodes, were infested. Of the 207 cladodes observed, 16 were damaged. The extent of infestation varied from 4.6\u0026ndash;11.9% (mean 7.7%). At Kundara, six cladodes out of twelve (50%) in a single clump, in a homestead, were observed affected. The infestation was also observed at Karingannur in Kollam on flowers and fruits, however, cladodes were free of ant damage. Besides cladode, flowers and fruits were also affected at Vembayam. At Maranallur, 45 out of 50 (90%) fruits and/ flowers on four infested clumps were damaged, however, no damage was observed on the cladodes.\u003c/p\u003e \u003cp\u003eAnts ate the areole (Figs.\u0026nbsp;4, 5) and the surrounding tissues at the base of spines, resulting in characteristic damage (Figs.\u0026nbsp;6, 7). Due to feeding of the tissues at the base of areole, deep emarginations were formed on the ridges of the cladode, in the place of spines (Fig.\u0026nbsp;6). The damaged portions turned yellow and then dark brown to black (Figs.\u0026nbsp;6, 7). The affected plants appeared weak and their growth was suppressed. At Vembayam, besides the base of spines, they also fed on flowers (Figs.\u0026nbsp;8, 9) and developing green fruits (Fig.\u0026nbsp;10) as well as mature, ripening fruits (Fig.\u0026nbsp;11). Ants fed directly on the flowers (Fig.\u0026nbsp;9) and fruits (Figs.\u0026nbsp;10, 11), resulting in sunken areas. On fruits, these sunken areas turned black (Figs.\u0026nbsp;10, 11). Infested fruits became visually unattractive and lost their market value.\u003c/p\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eIn Cactaceae, 156 species in 40 genera are known to have extrafloral nectaries (Keeler et al., 2015\u0026ndash;2024). \u003cem\u003eSelenicereus undatus\u003c/em\u003e (Haw.) D. R. Hunt, the white-fleshed pitahaya, possesses extrafloral nectaries (Keeler et al., 2015\u0026ndash;2024). It was observed that \u003cem\u003eS\u003c/em\u003e. \u003cem\u003ecostaricensis\u003c/em\u003e also have nectarines within the areole at the base of the spines as well as under the small, scale-like leaves beneath the areole on developing cladodes (Fig.\u0026nbsp;5). It is the extrafloral nectaries that attract \u003cem\u003eT. destructor\u003c/em\u003e to the dragon fruit cactus, which, according to Jerdon (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1851\u003c/span\u003e), \u0026lsquo;\u003cem\u003efeeds greedily on sugar\u0026rsquo;.\u003c/em\u003e\u003c/p\u003e \u003cp\u003eExtrafloral nectaries attract aggressive arthropods that protect developing leaves, shoots and flowers from herbivores and thus form an important weapon in the vast repertoire of plant defense mechanisms (Marazzi et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Heil, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). However, greater defense is accompanied by indirect costs such as reduced competitive ability, deterrence of pollinators (Ness, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) and natural enemies of herbivores (Callejas-Chavero et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) or exploitation of extrafloral nectaries by herbivores to their advantage when they are attracted to and retained by the extrafloral nectaries (Turlings \u0026amp; Wӓckers, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Popa, 2020). Selective, targeted feeding on extrafloral nectaries by several insect species, including some that are otherwise not primarily herbivorous, are known (Gish et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This is an extreme case of direct cost of production of extrafloral nectaries, which are supposed to be protective in function against herbivores. Here the \u0026lsquo;bodyguards\u0026rsquo; of the plant turn assailants, by shifting from predation to herbivory.\u003c/p\u003e \u003cp\u003eFlowers and fruits, when present, were preferred by the ants to the cladodes. This is probably due to the availability of more sugars and carbohydrates in fruits and flowers.\u003c/p\u003e \u003cp\u003eThis is the first report of \u003cem\u003eT. destructor\u003c/em\u003e as a pest of \u003cem\u003eS. costaracensis\u003c/em\u003e and the first report of extrafloral nectaries in \u003cem\u003eS. costaracensis\u003c/em\u003e.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAuthor contributions: AP independently collected data and made observations; ST conducted fieldwork, collected data and recorded observations; KDP conceptualized the study, carried out fieldwork, made observations and prepared the first draft of the manuscript. All authors reviewed and revised the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe are indebted to Sahana Shree, Ashoka Trust for Research in Ecology and the Environment (ATREE), Bengaluru for the identification of T. destructor. Sandra Maria Mathew prepared Figs 1 \u0026ndash; 3. This study was partially funded by the Kerala Agricultural University. Anooj S. S. critically reviewed the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAkheela, P., Santhoshkumar, T., \u0026amp; Varghese, T. S. (2024). First report of \u003cem\u003eBactrocera dorsalis\u003c/em\u003e (Hendel) (Diptera, Tephritidae) on the white-fleshed dragon fruit \u003cem\u003eSelenicereus undatus\u003c/em\u003e (Haworth) DR Hunt (Cactaceae) in India. \u003cem\u003eEntomon\u003c/em\u003e,\u0026nbsp;49(2), 283-286. https://doi.org/10.33307/entomon.v49i2.1190\u003c/li\u003e\n \u003cli\u003eBolton, B. (1987). A review of the \u003cem\u003eSolenopsis\u0026nbsp;\u003c/em\u003egenus-group and revision of Afrotropical \u003cem\u003eMonomorium\u0026nbsp;\u003c/em\u003eMayr (Hymenoptera: Formicidae). \u003cem\u003eBulletin of the British Museum (Natural History). Entomology\u0026nbsp;\u003c/em\u003e54(3), 263-452. https://biostor.org/reference/113868\u003c/li\u003e\n \u003cli\u003eCallejas-Chavero, A., Mart\u0026iacute;nez-Hern\u0026aacute;ndez, D., Flores-Mart\u0026iacute;nez, A., Moncada-Orellana, A., Diaz-Qui\u0026ntilde;ones, Y., \u0026amp; Vargas-Mendoza, C. F. (2020). Herbivory in Cacti: Fitness Effects of Two Herbivores, One Tending Ant on \u003cem\u003eMyrtillocactus geometrizans\u003c/em\u003e (Cactaceae). In: N\u0026uacute;\u0026ntilde;ez-Farf\u0026aacute;n, J., Valverde, P. (eds) Evolutionary Ecology of Plant-Herbivore Interaction. (pp. 109-134).\u0026nbsp;\u003cem\u003eSpringer, Cham\u003c/em\u003e.\u0026nbsp;\u003c/li\u003e\n \u003cli\u003eCollingwood, C. A., Tigar, B. J., \u0026amp; Agosti, D. (1997). Introduced ants in the United Arab Emirates. \u003cem\u003eJournal of Arid Environments\u003c/em\u003e 37(3), 505-512. https://doi.org/10.1006/jare.1997.0309\u003c/li\u003e\n \u003cli\u003eGish, M., Mescher, M. C., \u0026amp; DeMoraes, C. M. (2015). Targeted predation of extrafloral nectaries by insects despite localized chemical defences.\u0026nbsp;\u003cem\u003eProceedings of the Royal Society B: Biological Sciences\u0026nbsp;\u003c/em\u003e282(1816): 20151835. https://doi.org/10.1098/rspb.2015.1835\u003c/li\u003e\n \u003cli\u003eHarris, R., Abbott, K., Barton, K., Berry, J., Don, W., Gunawardana, D., Lester, P., Rees, J., Stanley M., Sutherland, A., \u0026amp; Toft, R. (2005). Invasive ant pest risk assessment project for Biosecurity New Zealand. Series of unpublished Landcare Research contract reports to Biosecurity New Zealand. BAH/35/2004-1.\u003c/li\u003e\n \u003cli\u003eHeil, M. (2015). Extrafloral nectar at the plant-insect interface: a spotlight on chemical ecology, phenotypic plasticity, and food webs. \u003cem\u003eAnnual Review of Entomology\u003c/em\u003e 60: 213-232.\u0026nbsp;https://doi.org/10.1146/annurev-ento-010814-020753\u003c/li\u003e\n \u003cli\u003eJerdon, T. C. (1851). A catalogue of the species of ants found in Southern India. \u003cem\u003eMadras Journal of Literature and Science\u003c/em\u003e 17, 103-127.\u003c/li\u003e\n \u003cli\u003eKavya, K. T. (2023). Management of phytophagous ants in vegetable crops under homestead cultivation. (Master\u0026rsquo;sThesis). Kerala Agricultural University, Vellayani. 83 p.\u003c/li\u003e\n \u003cli\u003eKeeler, K. H., Porturas, L. D., \u0026amp; Weber, M. G. (2015\u0026ndash;2024). World list of plants with extrafloral nectaries. \u0026nbsp;www.extrafloralnectaries.org. Accessed on 18 April 2024\u003c/li\u003e\n \u003cli\u003eMarazzi, B., Bronstein, J. L., \u0026amp; Koptur, S. (2013). The diversity, ecology and evolution of extrafloral nectaries: current perspectives and future challenges. \u003cem\u003eAnnals of Botany\u003c/em\u003e 111(6),1243-1250.\u0026nbsp;https://doi.org/10.1093/aob/mct109\u003c/li\u003e\n \u003cli\u003eNess, J. H. (2006). A mutualism\u0026apos;s indirect costs: the most aggressive plant bodyguards also deter pollinators. \u003cem\u003eOikos\u003c/em\u003e 113(3), 506-514. https://doi.org/10.1111/j.2006.0030-1299.14143.x\u003c/li\u003e\n \u003cli\u003ePopa, V. I., Georgescu, I.M., Potor, C. D., \u0026amp; Hoza, D. (2020). Defensive role of extrafloral nectaries. \u003cem\u003eScientific Papers. Series B. Horticulture\u0026nbsp;\u003c/em\u003e64(1),\u0026nbsp;693-700.\u003c/li\u003e\n \u003cli\u003ePrathapan, K. D., \u0026amp; Santhoshkumar, T. (2022). First report of \u003cem\u003eSpodoptera litura\u003c/em\u003e (Fabricius) (Noctuidae) on the dragon fruit species \u003cem\u003eSelenicereus costaricensis\u003c/em\u003e (F.A.C.Weber) S.Arias \u0026amp; N.Korotkova and \u003cem\u003eSelenicereus megalanthus\u003c/em\u003e (K.Schum. ex Vaupel) Moran (Cactaceae). \u003cem\u003eJournal of the Lepidopterists\u0026rsquo; Society\u003c/em\u003e 76 (4), 282-283.\u0026nbsp;https://doi.org/10.18473/lepi.76i4.a9\u003c/li\u003e\n \u003cli\u003eSharaf, M. R., Salman, S., Al Dhafer, H. M., Akbar, S. A., Abdel-Dayem, M. S., \u0026amp; Aldawood, A. S. (2016). Taxonomy and distribution of the genus \u003cem\u003eTrichomyrmex\u0026nbsp;\u003c/em\u003eMayr, 1865 (Hymenoptera: Formicidae) in the Arabian Peninsula, with the description of two new species\u003cem\u003e. European Journal of Taxonomy\u003c/em\u003e 246,1-36.\u0026nbsp;https://doi.org/10.5852/ejt.2016.246\u003c/li\u003e\n \u003cli\u003eTurlings, T. C. J., \u0026amp;Wӓckers, F. (2004). Recruitment of predators and parasitoids by herbivore-injured plants. In: \u003cem\u003eAdvances in Insect Chemical Ecology\u0026nbsp;\u003c/em\u003e(R. T. Card\u0026eacute;, ed.). (pp. 21-75). Cambridge University Press.\u0026nbsp;\u003c/li\u003e\n \u003cli\u003eWakchaure, G. C., Satish Kumar, Meena, K. K., Rane, J., \u0026amp; Pathak, H. (2021). Dragon Fruit Cultivation in India: Scope, Constraints and Policy Issues. \u003cem\u003eTechnical Bulletin No\u003c/em\u003e 27,47. ICAR\u0026ndash;National Institute of Abiotic Stress Management, Baramati, Pune, Maharashtra, India.\u003c/li\u003e\n \u003cli\u003eWay, M. J., Cammell, M. E., Bolton, B., \u0026amp; Kanagaratnam, P. (1989). Ants (Hymenoptera: Formicidae) as egg predators of coconut pests, especially in relation to biological control of the coconut caterpillar, \u003cem\u003eOpisina arenosella\u003c/em\u003e Walker (Lepidoptera: Xyloryctidae), in Sri Lanka. \u003cem\u003eBulletin of Entomological Research\u003c/em\u003e 79(2), 219-234.\u0026nbsp;https://doi.org/10.1017/S0007485300018204\u003c/li\u003e\n \u003cli\u003eWetterer, J. K. (2009). Worldwide spread of the destroyer ant, \u003cem\u003eMonomorium destructor\u003c/em\u003e (Hymenoptera: Formicidae). \u003cem\u003eMyrmecological News\u0026nbsp;\u003c/em\u003e12, 97-108.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Selenicereus costaricensis, India, Cactaceae, herbivory","lastPublishedDoi":"10.21203/rs.3.rs-4733344/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4733344/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe destroyer ant, \u003cem\u003eTrichomyrmex destructor\u003c/em\u003e (Jerdon, 1805) (Formicidae, Myrmicinae), is newly reported as a pest of the red dragon fruit \u003cem\u003eSelenicereus costaricensis\u003c/em\u003e (F. A. C. Weber) S. Arias \u0026amp; N. Korotkova ex Hammel (Cactaceae). It feeds on the areoles, scale leaves and surrounding green tissues at the base of spines of \u003cem\u003eS. costaricensis\u003c/em\u003e, due to the presence of extrafloral nectaries. The damaged portions turned yellow and then dark brown to black. The extent of infestation on the cladode was 0\u0026ndash;50%. Besides cladode, flowers and fruits were also damaged. Up to 90% fruits and/flowers were infested on four clumps. This is a case of direct cost of production of extrafloral nectaries, which are protective in function against herbivores. 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