Fruit-feeding butterfly diversity in a large city in southern India

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In a large city in southern India, we performed long-term sampling of fruit-feeding butterflies of two sites with mainly open habitats, and more modest sampling in two closed-canopy sites. We also sampled a nearby natural forest for comparison. We caught 3625 butterflies belonging to 29 species, of which sixteen species were recorded in the open habitats, and a further five species in the closed-canopy sites in the city. The open habitats in the city featured a protected species, the Gaudy Baron ( Euthalia lubentina (Cramer, 1777)), and closed-canopy sites had charismatic forest species such as the Clipper ( Parthenos sylvia (Cramer, [1775])) and Indian Nawab ( Polyura athamas (Drury) 1773). Community composition differed significantly among sites and sampling months. Overall, our results show that habitats within the city have fruit-feeding butterfly diversity comparable to that of natural forests, and are thus worth conserving for nature education. community composition host plant urban biodiversity protected species forest Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 INTRODUCTION Urban environments are home to a wide variety of wild animals and plants. This has two important consequences. Firstly, the parks, gardens, and wild corners in urban areas may conserve some species that do not thrive in the surrounding intensified agricultural landscapes (Kowarik 2011 ). Secondly, urban biodiversity may be the main contact that many people have with nature (Kowarik et al. 2025 ; Shwartz et al. 2014 ). While urban environments can, under some circumstances, serve as refuges for threatened or charismatic species, they are often less species-rich than natural habitats, as organisms face serious threats in urban environments (McKinney 2008 ). These include habitat loss and fragmentation, pollution (noise, light, chemicals, small particles), and a warmer climate (Aronson et al. 2017 ). Despite a growing interest in urban biodiversity, knowledge of biodiversity in tropical cities remains scarce (Newbold et al. 2020 ). It is estimated that urban areas cover over 2.3% of India's land area, and this percentage continues to increase (Pandey and Seto 2015). Currently, more than half of the people live and work in urban areas, and this is expected to increase to 65% in the coming decades (Sadashivam and Tabassu 2016 ), so that information on urban biodiversity becomes more critical. Studies on urban diversity in the region have found that considerable biodiversity can be found in gardens, forest fragments, and other habitats in cities (e. g. Mahesh et al. 2024 ; Vikas, Athreya, and Limaye 2022; Ravi and Thomas 2015; Kunhamu, Ajeesh, and Kumar 2015 ; Chowdhury et al. 2021 ; Rajashekara et al. 2025 ). Butterflies are a charismatic group that is often used in biodiversity studies and conservation outreach programs. Some tropical butterfly species in India show considerable adaptability by using anthropogenic resources as host plants for their larvae (Nitin et al. 2018 ; Kunte 2006 ; Kunte 2000 ; Rajashekara et al. 2025 ). For example, Tailed Palmflies ( Elymnias caudata Butler, 1871) use coconut palms, and Common Barons ( Euthalia aconthea (Cramer, [1777])) depend on mango trees (Nitin et al. 2018 ; Rajashekara et al. 2025 ). Adults of fruit-feeding species also use urban food sources by feeding on fallen cultivated fruits such as mangos, cashews, jackfruit, papaya, and bananas; and nectar-feeding species often visit cultivated, ornamental and exotic flowers (Rajashekara et al. 2025 ). These examples illustrate how some butterfly taxa can persist in modified landscapes. However, not all urban habitats will be equally suitable (e. g. Kuussaari et al. 2021; Dylewski, Maćkowiak, and Banaszak-Cibicka 2019 ; Rochat et al. 2017 ; Koh and Sodhi 2004; Chowdhury et al. 2017 ; Gupta, Tiwari, and Diwakar 2019; Pignataro et al. 2020 ; Ramírez-Restrepo and MacGregor-Fors 2017; Tam and Bonebrake 2016). In tropical forests, trapping fruit-feeding butterflies is a commonly used technique to assess biodiversity (e.g. DeVries, Hamm, and Fordyce 2016; Forsberg et al. 2020 ). It is also useful in more open tropical habitats (e.g. Freitas et al. 2021 ; Schmitt et al. 2021 ), and thus also holds promise for studies of biodiversity in tropical cities (Pignataro et al. 2020 ). This technique avoids observer bias and samples species that are otherwise rarely seen. Moreover, it allows for taking measurements and standardised photographs, and hands-on experience in nature-education settings, while no butterflies are harmed. We documented the community of fruit-feeding butterflies in two fragments of rural landscape (open habitats), and two closed-canopy sites within the city of Thiruvananthapuram, Kerala State, in southern India. We also sampled a natural semi-deciduous forest in Thiruvananthapuram District for comparison. METHODS Study site We sampled fruit-feeding butterflies in and near Thiruvananthapuram (Kerala State, southern India), a city with a metropolitan population estimated to be about 1.68 million as of March 2025. Its rapid urbanization has significant impacts on natural habitats (Chettry 2022 ; Cyriac 2022 ). At the city scale, from 2001 to 2021, urban areas have increased by 126%, while farmland and natural areas have decreased by 21%. Thus, wetlands and agricultural land have been lost, especially in the peripheral areas (Chettry 2022 ). We trapped butterflies in four sites in Thiruvananthapuram and one site near Vithura (Fig. 1 ). We sampled the Pongumoodu site (open habitat; 8°32’06.5”N, 76°54’53.9”E) for the longest period (Table 1 ). The site is along a small stream and is mainly comprised of abandoned rice paddy, which now features extensive coconut farming with grazing by cows and goats (Fig. 2 ). There are also some gardens with bananas, papaya, mango, and cashew. The Technopark site (open habitat; 8°32’52.4”N, 76°53’23.3”E), is next to the Technopark campus and is similar to the Pongumoodu site, but more extensive, with some open Eucalyptus forest, and open grassland and wetland grazed by cows, goats, and water buffalo (Fig. 2 ). Mookkunimala (closed canopy; 8°47’11.1”N, 77°01’16.3”E) is a rocky hill partly covered in rubber plantations, and steep and rocky sections are covered in wild thicket vegetation. Rubber trees are about 5–10 meters high, and the understory is regularly cut to allow easy access for rubber tapping and to reduce competition for the rubber trees, thus forming a forest-like habitat with a dense herb layer (Fig. 2 ). The site also features a stone quarry, and part of the hill is used as a shooting range (Fig. 1 ). Notably, our sampling was cut short here as bonnet macaques started to break our traps, as they eat both the bait and trapped insects. Church Forest (closed canopy; 8°33′40.3″N 76°54′29.9″E), is a small forested valley owned by a church (Fig. 2 ), with some coconut and old fruit trees (mango, cashew), with on higher ground a small rubber plantation. The Vithura site (natural forest; 8°41'01.3"N, 77°08'18.5"E) is a natural semi-deciduous forest within the campus of the Indian Institute for Science Education and Research, Thiruvananthapuram (Fig. 1 ). It borders the protected forest of the Peppara Wildlife Sanctuary in Agasthyamalai Biosphere Reserve. We also sampled a rubber plantation next to the forest. Table 1 Habitat of sampling sites, sampling period, and the number of traps deployed at each site. Site Site Type Habitat Sampling Period # Traps Pongumoodu Open coconut, banana, grazing May 2016–March 2018 12 Technopark Open coconut, banana, grazing Nov 2016–March 2018 15 Mookkunimala Closed-canopy rubber, wild bush July 2016–Nov 2016 15 Church forest Closed-canopy abandoned orchard, rubber Dec 2016–May 2017 10 Vithura Natural forest semi-deciduous forest, rubber Jun 2016–Jan 2017 30 Butterfly Sampling At each site, we caught butterflies with fruit-baited traps (Fig. 2 ). The bait consisted of bananas that were sliced and mashed and left to ferment for at least three days in a loosely closed bucket. At the two open-habitat sites, we usually baited traps every two weeks with a serving spoonful of bait, and we then typically serviced the traps during the subsequent 2–3 days (Fig. 3 ), usually during the early afternoon (2–3 PM), adding bait when necessary. The other sites were sampled more sporadically (Fig. 3 ). We photographed the butterflies before release. We identified the butterflies on the photographs with the help of the iFoundButterflies website (Kunte, Sondhi, and Roy 2025 ) and Kunte ( 2000 ). Data Analysis To visually assess the degree of sample completeness, we generated rarefaction curves for each site. To visualise how community composition changed over time at each site, we made a non-metric multidimensional scaling (NMDS) plot. Specifically, we first pooled all observations by month-year combination for each site (months often including two sampling campaigns). We then used NMDS to plot communities that were more similar closer together in a two-dimensional space. To visualise the temporal change in community composition for each site, we connected communities from subsequent months with arrows and labelled points with month and year. To statistically test for site and temporal effects, we performed PERMANOVA analysis. We performed all analyses in R (R_Core_Team 2025) using the package vegan (Oksanen et al. 2007 ), and drew plots using the ggplot2 package (Wickham 2016). RESULTS Overall, we caught 3625 butterflies belonging to 29 species in the urban sites and natural forest combined (Table 2 ). The most common species was the Common Evening Brown ( Melanitis leda (L. 1758); Fig. 4 ), followed by the Dark-branded Bushbrown ( Mycalesis mineus (L. 1758)), and the Chocolate Pansy ( Junonia iphita (Cramer, [1779]); Table 2 ). Table 2 The number of individuals per species and site sorted by overall abundance. Pongumoodu and Technopark had mainly open habitats, while Church Forest and Mookkunimala were mainly closed-canopy sites within the city, and Vithura was the natural forest site. Species Common name Pongumoodu Technopark Church Forest Mookkunimala Vithura Total Melanitis leda (Linnaeus, 1758) Common Evening Brown 587 629 13 163 74 1466 Mycalesis mineus (Linnaeus, 1758) Dark-branded Bushbrown 346 201 8 12 13 580 Junonia iphita (Cramer, [1779]) Chocolate Pansy 102 115 10 212 7 446 Ariadne ariadne (Linnaeus, 1763) Angled Castor 102 148 15 24 3 292 Tanaecia lepidea (Butler, 1868) Grey Count 57 98 6 6 13 180 Euthalia aconthea (Cramer, [1777]) Common Baron 63 73 13 13 2 164 Elymnias caudata Butler, 1871 Tailed Palmfly 73 73 1 1 4 152 Ariadne merione (Cramer, [1777]) Common Castor 28 67 2 9 0 106 Hypolimnas bolina (Linnaeus, 1758) Great Eggfly 7 13 1 24 4 49 Charaxes solon (Fabricius, 1793) Black Rajah 13 5 1 18 0 37 Junonia atlites (Linnaeus, 1763) Grey Pansy 19 6 0 1 0 26 Lethe drypetis (Hewitson, 1863) Tamil Treebrown 1 0 0 6 15 22 Mycalesis patnia (Moore, 1857) Gladeye Bushbrown 0 0 0 16 3 19 Melanitis phedima (Cramer, [1780]) Dark Evening Brown 0 0 1 3 11 15 Orsotriaena medus (Fabricius, 1775) Medus Brown 6 1 1 2 1 11 Polyura athamas (Drury) 1773 Indian Nawab 0 0 2 9 0 11 Kaniska canace (Linnaeus, 1763) Blue Admiral 0 1 0 3 6 10 Lethe europa (Fabricius, 1787) Bamboo Treebrown 0 0 0 5 4 9 Euthalia lubentina (Cramer, 1777) Gaudy Baron 2 2 0 2 0 6 Junonia lemonias (Linnaeus, 1758) Lemon Pansy 1 4 0 0 0 5 Melanitis zitenius (Herbst, 1796) Great Evening Brown 0 0 0 0 4 4 Moduza procris (Cramer, [1777]) Commander 0 3 0 0 1 4 Charaxes bernardus (Fabricius, 1793) Tawny Rajah 0 0 0 1 1 2 Lethe rohria (Fabricius, 1787) Common Treebrown 0 0 0 0 2 2 Parthenos sylvia (Cramer, [1775]) Clipper 0 0 2 0 0 2 Zipaetis saitis Hewitson, [1863] Banded Catseye 0 0 0 0 2 2 Amathusia travancorica Kunte, 2024 Travancore Palmking 0 0 0 0 1 1 Mycalesis anaxias Hewitson, 1862 White-bar Bushbrown 0 0 0 0 1 1 Parantirrhoea marshalli Wood-Mason, [1881] Travancore Evening Brown 0 0 0 0 1 1 The rarefaction curve for samples from the Pongumoodu site (open habitat) showed that sampling completeness was high (Fig. 5 ), with diversity confidently estimated at 15 species. Thus, further sampling is unlikely to recover a substantial number of additional species. Similarly, the rarefaction curve for Technopark (open habitat) showed high sample completeness (Fig. 5 ), with diversity estimated at 16 species. The number of species for these open-habitat sites combined was 17, and included the Gaudy Baron ( Euthalia lubentina (Cramer, 1777); Fig. 4 ) in both. The species composition of these two sites was very similar, with one Tamil Tree-brown ( Lethe drypetis (Hewitson, 1863)) trapped in Pongumoodu and one Blue Admiral ( Kaniska canace (Linnaeus, 1763)) and three Commanders ( Moduza procris (Cramer, [1777])) in the Technopark site as the exceptions (Table 2 ). At the Church Forest site (closed canopy habitat), the butterfly community remained under-sampled (Fig. 5 ). The steep rarefaction curve indicates a quite high diversity. In particular, this is the only site where the Clipper ( Parthenos sylvia (Cramer, [1775])), a charismatic species associated with forests, was found (Table 2 , Fig. 4 ). Sampling at Mookkunimala (closed canopy habitat) also resulted in fairly adequate sample completeness (Fig. 5 ). Nevertheless, the curve indicates that a few more species than the 20 found would have been sampled if sampling there had not been cut short by the monkeys. The sampling at Vithura was also incomplete (Fig. 5 ), but butterfly diversity was highest, including characteristic forest species like the Large Evening Brown ( Me. Zitenius (Herbst, 1796)), the newly recognised Travancore Palmking (Amathusia travancorica Kunte 2024 ; Kunte and Basu 2024 ), and the enigmatic Travancore Evening Brown ( Parantirrhoea marshalli Wood-Mason, [1881]; Table 2 ). The NMDS plot shows that the extent of temporal change in community composition obscures differences between sites. Nevertheless, the most natural site, Vithura, is situated on the margin of the plot (Fig. 6 ), as it features several species not found in other sites (Table 2 ). Overall, the differences between sites in community composition were significant (Table 3 ). Table 3 Results of PERMANOVA analysis, testing for effects of site and month-year on butterfly community composition. SoS = sum of squares. Df SoS R 2 F-value P-value Site 4 1.438 0.166 2.518 0.001 Month*Year 20 5.062 0.586 1.773 0.003 Residual 15 2.142 0.248 Total 39 8.642 1.000 DISCUSSION We sampled fruit-feeding butterflies in Thiruvananthapuram and a nearby natural forest, and showed that green areas in the city harbour significant fruit-feeding butterfly faunas, albeit impoverished compared to natural sites. Some forest species managed to persist in a forest fragment, and even occurred in rubber plantations. Notably, we found Clippers ( P. sylvia ) in the small Church Forest. We also found a rich fauna on the large hill with rubber plantations and wild thicket vegetation (Mookkunimala). In the open-habitat sites (Pongumoodu and Technopark), we found a more modest species richness despite higher sampling intensity. Nevertheless, there were some interesting species there too, particularly the Gaudy Baron ( Eu. lubentina ), which is protected in India under Schedule IV of the Wildlife (Protection) Act, 1972, and was not encountered in natural forests (Das, Radhamany, and Molleman 2022). Notably, we do not have good natural controls to compare our urban sites with. Without human influence, the area where Thiruvananthapuram city is now situated would have been mainly lowland rainforest, a vegetation type that is almost non-existent in this region today. The nearest natural vegetation available to us was semi-deciduous forest at the foot of the Western Ghats. However, this may be similar to what would cover Mookkunimala if there had been no human influence. A year-long study in Silent Valley National Park, a mid-altitude rainforest further north in the Western Ghats, found 32 species across forest interior and edge habitats (Das, Radhamany, and Molleman 2022). Most of the species we caught were also found there, except that we observed Travancore Palmking ( A. phidippus ), Black Rajah ( Charaxes solon ), Tailed Palmfly ( El. Caudata ), Gaudy Baron ( Eu. lubentina ), Lemon Pansy ( Junonia lemonias (Linnaeus, 1758)), Blue Admiral ( K. canace ), Tamil Treebrown ( Lethe drypetis (Hewitson, 1863)), Tranvancore Evening Brown ( P. marshalli ), and Common Nawab ( P. athamas ), not observed in that study. Overall, we found that in the open habitat sites, we see a subset of the species found in more natural sites in the region, most of which are more typical for open habitats, but a few forest species persist as well. The lower species richness of fruit-feeding butterflies in more open habitats may reflect that this guild is generally more diverse in forests than in more open vegetation (Schmitt et al. 2021 ). The butterfly species composition at our sites could in part be attributed to host plants. The common species at our open-habitat sites tended to be associated with either grasses (e.g. Common Evening Brown ( Me. leda ) and Black-branded Bushbrown ( My. mineus )), cultivated trees (e.g. Common Baron ( Eu. aconthea ), Tailed Palmfly ( El. caudata )), or ruderal plants (e.g. Castors and Pansies ( Ariadne and Junonia species); Nitin et al. 2018 ). The Gaudy Baron ( Eu. lubentina ) is associated with mistletoes ( Dendrophthoe sp.). These mistletoes are often growing on mango, fig, and tamarind, trees that are common in the city as fruit or shade trees (Ravi and Thomas 2015). Thus, conserving these vegetations is an important pillar for conserving butterflies in urban regions. There are at least two takeaways from this study. Firstly, these green areas within the urban matrix do support significant butterfly diversity that can also be exploited for nature education. For example, pupils could perform butterfly trapping on school campuses or during excursions on foot. Secondly, the temporal variation in community composition shows that to compare the butterfly diversity across urban sites, they have to be sampled over extended periods. This cannot simply be avoided by sampling for a short time in the different sites simultaneously because temporal patterns may differ among sites. The rarefaction curves indicate that a suitable sample size for quantifying differences in butterfly community parameters between sites in this system would be about 500 individuals per site. Declarations Author Contribution FM performed the sampling and supervised BCA, followed by data validation, final data analysis, and writing the final version of the manuscript. BCA performed species identification and data basing, initial data analysis, and wrote the initial version of the manuscript. Acknowledgement We thank Ullasa Kodandaramaiah for supporting the fieldwork, and Kalesh Sadasivan for advice on sampling sites, identifications, and corrections on the manuscript. For help with the fieldwork, we are grateful to Sridhar Halali, Dheeraj Halali, Jisha Vijaykumar, O. Vignesh, Bharat Parthasarathy, and Soumen Mallick. The fieldwork of FM was funded through an INSPIRE Faculty Award to Ullasa Kodandaramaiah (DST/INSPIRE/04/2013/000476), and data curation to manuscript preparation was made possible by grant 2021/43/B/NZ8/00966 from the Naradowe Centrum Nauki (National Science Centre, Poland). Data Availability Data is provided within the supplementary information file References Aronson, Myla FJ, Christopher A Lepczyk, Karl L Evans, Mark A Goddard, Susannah B Lerman, J Scott MacIvor, Charles H Nilon, and Timothy Vargo. 2017. 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Supplementary Files TVMbutterflyMonitoringUrbanEcologies.xlsx Cite Share Download PDF Status: Published Journal Publication published 19 Dec, 2025 Read the published version in Urban Ecosystems → Version 1 posted Editorial decision: Revision requested 07 Nov, 2025 Reviews received at journal 06 Nov, 2025 Reviews received at journal 04 Nov, 2025 Reviews received at journal 17 Oct, 2025 Reviewers agreed at journal 16 Oct, 2025 Reviewers agreed at journal 16 Oct, 2025 Reviews received at journal 16 Oct, 2025 Reviewers agreed at journal 16 Oct, 2025 Reviewers agreed at journal 15 Oct, 2025 Reviewers agreed at journal 15 Oct, 2025 Reviewers agreed at journal 15 Oct, 2025 Reviewers invited by journal 14 Oct, 2025 Editor assigned by journal 08 Oct, 2025 Submission checks completed at journal 06 Oct, 2025 First submitted to journal 29 Sep, 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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Molleman","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAv0lEQVRIiWNgGAWjYBAC9gYQWWAD4fEAMR8hLTwHQKRBGkILG5FaDpOiRfrw0Q0fDM7L685IYHzwto0hj7AWvrS0mzMMbhtuu5HAbDi3jaGYoBZ7Hh6z2zwGtxmBWtikedsYEtsI2sLD/w2o5Zw9UAv7byK18LABtRxIBNnCTKQWNjOgX5KTt5152Cw555wEYb/w8DA/u/Ghws522/Hkgx/elNnk8RPSggQYG4CERAIJOqCADC2jYBSMglEw3AEAxLo6O137QrwAAAAASUVORK5CYII=","orcid":"","institution":"Adam Mickiewicz University Poznań","correspondingAuthor":true,"prefix":"","firstName":"Freerk","middleName":"","lastName":"Molleman","suffix":""},{"id":534696188,"identity":"b87f91bd-198a-4434-ba1c-b7ad1becd104","order_by":1,"name":"Berke Can Aydin","email":"","orcid":"","institution":"Afyon Kocatepe University","correspondingAuthor":false,"prefix":"","firstName":"Berke","middleName":"Can","lastName":"Aydin","suffix":""}],"badges":[],"createdAt":"2025-09-29 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19:25:18","extension":"png","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":25544,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/66cb12e9da1fcd39f79d1245.png"},{"id":94700137,"identity":"24baa3dd-8775-41cf-9163-dd60b80baadf","added_by":"auto","created_at":"2025-10-29 19:25:18","extension":"xml","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":103316,"visible":true,"origin":"","legend":"","description":"","filename":"2ad3cbd8adfb445b8f47caf077d57ebc1structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/94ea7f231ee4394e692f47eb.xml"},{"id":94728291,"identity":"f75870f3-95ea-4628-9cba-ed5132b2a6d8","added_by":"auto","created_at":"2025-10-30 07:03:28","extension":"html","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":110546,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/e4464945769a87a4fcfa1d48.html"},{"id":94700111,"identity":"88c66933-4b35-4471-b28b-189d46a52501","added_by":"auto","created_at":"2025-10-29 19:25:17","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":697612,"visible":true,"origin":"","legend":"\u003cp\u003eSatellite photos of the study sites (Google Earth, © 2025 Airbus and Maxar Technologies).\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/4c0c4e6fa8a7b9c670cdbd0b.png"},{"id":94700130,"identity":"03da60bd-567e-447b-92e2-0626739606b2","added_by":"auto","created_at":"2025-10-29 19:25:17","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":715746,"visible":true,"origin":"","legend":"\u003cp\u003eHabitat photos of the urban study sites (photos: FM). Note that each site was heterogeneous, including various habitat types. A butterfly trap is clearly visible in the photograph from Church Forest.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/89b662f2f4ec13da4c1d8439.png"},{"id":94700118,"identity":"7f7a2d20-5fda-467d-ace5-9067841c3988","added_by":"auto","created_at":"2025-10-29 19:25:17","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":104999,"visible":true,"origin":"","legend":"\u003cp\u003eTiming of butterfly sampling for each site with the log abundance of butterflies caught at a given day. Graphs are stacked for comparison. There was always something caught on each trapping day.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/8fb25c94c0a7d0c82e46e48f.png"},{"id":94729591,"identity":"e245fad4-6f92-4054-a086-05a5700c6ba2","added_by":"auto","created_at":"2025-10-30 07:05:11","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":522401,"visible":true,"origin":"","legend":"\u003cp\u003eThe most common species (Common Evening Brown, \u003cem\u003eMelanitis leda\u003c/em\u003e (Linnaeus, 1758)), the largest species (Clipper, \u003cem\u003eParthenos sylvia\u003c/em\u003e (Cramer, [1775])), and the protected species (Gaudy Baron, \u003cem\u003eEuthalia lubentina \u003c/em\u003e(Cramer, 1777)), found within the urban sites (photos: FM). Butterflies were photographed and released.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/e091eb9a414cb3a15dfdb2e3.png"},{"id":94700115,"identity":"8f1e7480-bc41-44ae-873b-0507ef55fb7b","added_by":"auto","created_at":"2025-10-29 19:25:17","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":167562,"visible":true,"origin":"","legend":"\u003cp\u003eRarefaction curves for each sampled site, where levelling off indicates high sample completeness, and high initial steepness indicates high species diversity.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/6655dcb69777e645ba75396a.png"},{"id":94729462,"identity":"cdc2985f-ea5a-48fc-83aa-52216851d8e3","added_by":"auto","created_at":"2025-10-30 07:04:59","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":167300,"visible":true,"origin":"","legend":"\u003cp\u003eNon-metric multidimensional scaling results of butterfly communities by month and site, where consecutive months are connected by arrows for each site. When points are near each other, the butterfly communities were similar.\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/64904f9082afaa04d87111a2.png"},{"id":98813836,"identity":"11621a84-14ed-4cb0-ad8e-4a3d75dcdf86","added_by":"auto","created_at":"2025-12-22 16:04:20","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3087399,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/75ae9acb-f6f9-4bf3-aba5-e04ee69ce721.pdf"},{"id":94700114,"identity":"37a17b03-f301-4871-ba42-4ed7405e0bc6","added_by":"auto","created_at":"2025-10-29 19:25:17","extension":"xlsx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":174835,"visible":true,"origin":"","legend":"","description":"","filename":"TVMbutterflyMonitoringUrbanEcologies.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7740235/v1/f5d0ea8c95fe86c590ef3163.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Fruit-feeding butterfly diversity in a large city in southern India","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eUrban environments are home to a wide variety of wild animals and plants. This has two important consequences. Firstly, the parks, gardens, and wild corners in urban areas may conserve some species that do not thrive in the surrounding intensified agricultural landscapes (Kowarik \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Secondly, urban biodiversity may be the main contact that many people have with nature (Kowarik et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Shwartz et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). While urban environments can, under some circumstances, serve as refuges for threatened or charismatic species, they are often less species-rich than natural habitats, as organisms face serious threats in urban environments (McKinney \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). These include habitat loss and fragmentation, pollution (noise, light, chemicals, small particles), and a warmer climate (Aronson et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Despite a growing interest in urban biodiversity, knowledge of biodiversity in tropical cities remains scarce (Newbold et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIt is estimated that urban areas cover over 2.3% of India's land area, and this percentage continues to increase (Pandey and Seto 2015). Currently, more than half of the people live and work in urban areas, and this is expected to increase to 65% in the coming decades (Sadashivam and Tabassu \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), so that information on urban biodiversity becomes more critical. Studies on urban diversity in the region have found that considerable biodiversity can be found in gardens, forest fragments, and other habitats in cities (e. g. Mahesh et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Vikas, Athreya, and Limaye 2022; Ravi and Thomas 2015; Kunhamu, Ajeesh, and Kumar \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Chowdhury et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rajashekara et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eButterflies are a charismatic group that is often used in biodiversity studies and conservation outreach programs. Some tropical butterfly species in India show considerable adaptability by using anthropogenic resources as host plants for their larvae (Nitin et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Kunte \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Kunte \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Rajashekara et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For example, Tailed Palmflies (\u003cem\u003eElymnias caudata\u003c/em\u003e Butler, 1871) use coconut palms, and Common Barons (\u003cem\u003eEuthalia aconthea\u003c/em\u003e (Cramer, [1777])) depend on mango trees (Nitin et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Rajashekara et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Adults of fruit-feeding species also use urban food sources by feeding on fallen cultivated fruits such as mangos, cashews, jackfruit, papaya, and bananas; and nectar-feeding species often visit cultivated, ornamental and exotic flowers (Rajashekara et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These examples illustrate how some butterfly taxa can persist in modified landscapes. However, not all urban habitats will be equally suitable (e. g. Kuussaari et al. 2021; Dylewski, Maćkowiak, and Banaszak-Cibicka \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Rochat et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Koh and Sodhi 2004; Chowdhury et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Gupta, Tiwari, and Diwakar 2019; Pignataro et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ram\u0026iacute;rez-Restrepo and MacGregor-Fors 2017; Tam and Bonebrake 2016).\u003c/p\u003e\u003cp\u003eIn tropical forests, trapping fruit-feeding butterflies is a commonly used technique to assess biodiversity (e.g. DeVries, Hamm, and Fordyce 2016; Forsberg et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). It is also useful in more open tropical habitats (e.g. Freitas et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Schmitt et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), and thus also holds promise for studies of biodiversity in tropical cities (Pignataro et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This technique avoids observer bias and samples species that are otherwise rarely seen. Moreover, it allows for taking measurements and standardised photographs, and hands-on experience in nature-education settings, while no butterflies are harmed.\u003c/p\u003e\u003cp\u003eWe documented the community of fruit-feeding butterflies in two fragments of rural landscape (open habitats), and two closed-canopy sites within the city of Thiruvananthapuram, Kerala State, in southern India. We also sampled a natural semi-deciduous forest in Thiruvananthapuram District for comparison.\u003c/p\u003e"},{"header":"METHODS","content":"\u003cp\u003e\u003cstrong\u003eStudy site\u003c/strong\u003e\u003cp\u003eWe sampled fruit-feeding butterflies in and near Thiruvananthapuram (Kerala State, southern India), a city with a metropolitan population estimated to be about 1.68\u0026nbsp;million as of March 2025. Its rapid urbanization has significant impacts on natural habitats (Chettry \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Cyriac \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). At the city scale, from 2001 to 2021, urban areas have increased by 126%, while farmland and natural areas have decreased by 21%. Thus, wetlands and agricultural land have been lost, especially in the peripheral areas (Chettry \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003c/p\u003e\u003cp\u003eWe trapped butterflies in four sites in Thiruvananthapuram and one site near Vithura (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We sampled the Pongumoodu site (open habitat; 8\u0026deg;32\u0026rsquo;06.5\u0026rdquo;N, 76\u0026deg;54\u0026rsquo;53.9\u0026rdquo;E) for the longest period (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The site is along a small stream and is mainly comprised of abandoned rice paddy, which now features extensive coconut farming with grazing by cows and goats (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). There are also some gardens with bananas, papaya, mango, and cashew. The Technopark site (open habitat; 8\u0026deg;32\u0026rsquo;52.4\u0026rdquo;N, 76\u0026deg;53\u0026rsquo;23.3\u0026rdquo;E), is next to the Technopark campus and is similar to the Pongumoodu site, but more extensive, with some open Eucalyptus forest, and open grassland and wetland grazed by cows, goats, and water buffalo (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Mookkunimala (closed canopy; 8\u0026deg;47\u0026rsquo;11.1\u0026rdquo;N, 77\u0026deg;01\u0026rsquo;16.3\u0026rdquo;E) is a rocky hill partly covered in rubber plantations, and steep and rocky sections are covered in wild thicket vegetation. Rubber trees are about 5\u0026ndash;10 meters high, and the understory is regularly cut to allow easy access for rubber tapping and to reduce competition for the rubber trees, thus forming a forest-like habitat with a dense herb layer (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The site also features a stone quarry, and part of the hill is used as a shooting range (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Notably, our sampling was cut short here as bonnet macaques started to break our traps, as they eat both the bait and trapped insects. Church Forest (closed canopy; 8\u0026deg;33\u0026prime;40.3\u0026Prime;N 76\u0026deg;54\u0026prime;29.9\u0026Prime;E), is a small forested valley owned by a church (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), with some coconut and old fruit trees (mango, cashew), with on higher ground a small rubber plantation. The Vithura site (natural forest; 8\u0026deg;41'01.3\"N, 77\u0026deg;08'18.5\"E) is a natural semi-deciduous forest within the campus of the Indian Institute for Science Education and Research, Thiruvananthapuram (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). It borders the protected forest of the Peppara Wildlife Sanctuary in Agasthyamalai Biosphere Reserve. We also sampled a rubber plantation next to the forest.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\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\u003eHabitat of sampling sites, sampling period, and the number of traps deployed at each site.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSite\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSite Type\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eHabitat\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSampling Period\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003e# Traps\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePongumoodu\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eOpen\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ecoconut, banana, grazing\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eMay 2016\u0026ndash;March 2018\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTechnopark\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eOpen\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ecoconut, banana, grazing\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eNov 2016\u0026ndash;March 2018\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMookkunimala\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eClosed-canopy\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003erubber, wild bush\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eJuly 2016\u0026ndash;Nov 2016\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eChurch forest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eClosed-canopy\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eabandoned orchard, rubber\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eDec 2016\u0026ndash;May 2017\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVithura\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNatural forest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003esemi-deciduous forest, rubber\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e\u003cp\u003eJun 2016\u0026ndash;Jan 2017\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eButterfly Sampling\u003c/strong\u003e\u003cp\u003eAt each site, we caught butterflies with fruit-baited traps (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The bait consisted of bananas that were sliced and mashed and left to ferment for at least three days in a loosely closed bucket. At the two open-habitat sites, we usually baited traps every two weeks with a serving spoonful of bait, and we then typically serviced the traps during the subsequent 2\u0026ndash;3 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), usually during the early afternoon (2\u0026ndash;3 PM), adding bait when necessary. The other sites were sampled more sporadically (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). We photographed the butterflies before release. We identified the butterflies on the photographs with the help of the iFoundButterflies website (Kunte, Sondhi, and Roy \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and Kunte (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eData Analysis\u003c/strong\u003e\u003cp\u003eTo visually assess the degree of sample completeness, we generated rarefaction curves for each site. To visualise how community composition changed over time at each site, we made a non-metric multidimensional scaling (NMDS) plot. Specifically, we first pooled all observations by month-year combination for each site (months often including two sampling campaigns). We then used NMDS to plot communities that were more similar closer together in a two-dimensional space. To visualise the temporal change in community composition for each site, we connected communities from subsequent months with arrows and labelled points with month and year. To statistically test for site and temporal effects, we performed PERMANOVA analysis. We performed all analyses in R (R_Core_Team 2025) using the package vegan (Oksanen et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), and drew plots using the ggplot2 package (Wickham 2016).\u003c/p\u003e\u003c/p\u003e"},{"header":"RESULTS","content":"\u003cp\u003eOverall, we caught 3625 butterflies belonging to 29 species in the urban sites and natural forest combined (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The most common species was the Common Evening Brown (\u003cem\u003eMelanitis leda\u003c/em\u003e (L. 1758); Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), followed by the Dark-branded Bushbrown (\u003cem\u003eMycalesis mineus\u003c/em\u003e (L. 1758)), and the Chocolate Pansy (\u003cem\u003eJunonia iphita\u003c/em\u003e (Cramer, [1779]); Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThe number of individuals per species and site sorted by overall abundance. Pongumoodu and Technopark had mainly open habitats, while Church Forest and Mookkunimala were mainly closed-canopy sites within the city, and Vithura was the natural forest site.\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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommon name\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePongumoodu\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eTechnopark\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChurch Forest\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eMookkunimala\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eVithura\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eTotal\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMelanitis leda\u003c/em\u003e (Linnaeus, 1758)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommon Evening Brown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e587\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e629\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e163\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e74\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1466\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMycalesis mineus\u003c/em\u003e (Linnaeus, 1758)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDark-branded Bushbrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e346\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e201\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e580\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eJunonia iphita\u003c/em\u003e (Cramer, [1779])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChocolate Pansy\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e102\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e115\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e212\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e446\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAriadne ariadne\u003c/em\u003e (Linnaeus, 1763)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAngled Castor\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e102\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e148\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e292\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eTanaecia lepidea\u003c/em\u003e (Butler, 1868)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGrey Count\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e57\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e98\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6\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\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e180\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eEuthalia aconthea\u003c/em\u003e (Cramer, [1777])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommon Baron\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e63\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e164\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eElymnias caudata\u003c/em\u003e Butler, 1871\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTailed Palmfly\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\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\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e152\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAriadne merione\u003c/em\u003e (Cramer, [1777])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommon Castor\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e67\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9\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\u003e106\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eHypolimnas bolina\u003c/em\u003e (Linnaeus, 1758)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreat Eggfly\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e49\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCharaxes solon\u003c/em\u003e (Fabricius, 1793)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eBlack Rajah\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e18\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\u003e37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eJunonia atlites\u003c/em\u003e (Linnaeus, 1763)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGrey Pansy\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\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\u003e26\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLethe drypetis\u003c/em\u003e (Hewitson, 1863)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTamil Treebrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\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\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMycalesis patnia\u003c/em\u003e (Moore, 1857)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGladeye Bushbrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMelanitis phedima\u003c/em\u003e (Cramer, [1780])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDark Evening Brown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eOrsotriaena medus\u003c/em\u003e (Fabricius, 1775)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMedus Brown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003ePolyura athamas\u003c/em\u003e (Drury) 1773\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eIndian Nawab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e9\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\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eKaniska canace\u003c/em\u003e (Linnaeus, 1763)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eBlue Admiral\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLethe europa\u003c/em\u003e (Fabricius, 1787)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eBamboo Treebrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eEuthalia lubentina\u003c/em\u003e (Cramer, 1777)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGaudy Baron\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e2\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\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eJunonia lemonias\u003c/em\u003e (Linnaeus, 1758)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLemon Pansy\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\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\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMelanitis zitenius\u003c/em\u003e (Herbst, 1796)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGreat Evening Brown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eModuza procris\u003c/em\u003e (Cramer, [1777])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommander\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCharaxes bernardus\u003c/em\u003e (Fabricius, 1793)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTawny Rajah\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLethe rohria\u003c/em\u003e (Fabricius, 1787)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCommon Treebrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eParthenos sylvia\u003c/em\u003e (Cramer, [1775])\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eClipper\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\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\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eZipaetis saitis\u003c/em\u003e Hewitson, [1863]\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eBanded Catseye\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAmathusia travancorica\u003c/em\u003e Kunte, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTravancore Palmking\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eMycalesis anaxias\u003c/em\u003e Hewitson, 1862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eWhite-bar Bushbrown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eParantirrhoea marshalli\u003c/em\u003e Wood-Mason, [1881]\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTravancore Evening Brown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThe rarefaction curve for samples from the Pongumoodu site (open habitat) showed that sampling completeness was high (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), with diversity confidently estimated at 15 species. Thus, further sampling is unlikely to recover a substantial number of additional species. Similarly, the rarefaction curve for Technopark (open habitat) showed high sample completeness (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), with diversity estimated at 16 species. The number of species for these open-habitat sites combined was 17, and included the Gaudy Baron (\u003cem\u003eEuthalia lubentina\u003c/em\u003e (Cramer, 1777); Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e) in both. The species composition of these two sites was very similar, with one Tamil Tree-brown (\u003cem\u003eLethe drypetis\u003c/em\u003e (Hewitson, 1863)) trapped in Pongumoodu and one Blue Admiral (\u003cem\u003eKaniska canace\u003c/em\u003e (Linnaeus, 1763)) and three Commanders (\u003cem\u003eModuza procris\u003c/em\u003e (Cramer, [1777])) in the Technopark site as the exceptions (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). At the Church Forest site (closed canopy habitat), the butterfly community remained under-sampled (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). The steep rarefaction curve indicates a quite high diversity. In particular, this is the only site where the Clipper (\u003cem\u003eParthenos sylvia\u003c/em\u003e (Cramer, [1775])), a charismatic species associated with forests, was found (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Sampling at Mookkunimala (closed canopy habitat) also resulted in fairly adequate sample completeness (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Nevertheless, the curve indicates that a few more species than the 20 found would have been sampled if sampling there had not been cut short by the monkeys. The sampling at Vithura was also incomplete (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), but butterfly diversity was highest, including characteristic forest species like the Large Evening Brown (\u003cem\u003eMe. Zitenius\u003c/em\u003e (Herbst, 1796)), the newly recognised Travancore Palmking (Amathusia travancorica Kunte \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Kunte and Basu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and the enigmatic Travancore Evening Brown (\u003cem\u003eParantirrhoea marshalli\u003c/em\u003e Wood-Mason, [1881]; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe NMDS plot shows that the extent of temporal change in community composition obscures differences between sites. Nevertheless, the most natural site, Vithura, is situated on the margin of the plot (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e), as it features several species not found in other sites (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Overall, the differences between sites in community composition were significant (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eResults of PERMANOVA analysis, testing for effects of site and month-year on butterfly community composition. SoS\u0026thinsp;=\u0026thinsp;sum of squares.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDf\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSoS\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eR\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eF-value\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eP-value\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSite\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.438\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.166\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.518\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMonth*Year\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.062\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.586\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.773\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.003\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eResidual\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.142\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.248\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTotal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e39\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e8.642\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eWe sampled fruit-feeding butterflies in Thiruvananthapuram and a nearby natural forest, and showed that green areas in the city harbour significant fruit-feeding butterfly faunas, albeit impoverished compared to natural sites. Some forest species managed to persist in a forest fragment, and even occurred in rubber plantations. Notably, we found Clippers (\u003cem\u003eP. sylvia\u003c/em\u003e) in the small Church Forest. We also found a rich fauna on the large hill with rubber plantations and wild thicket vegetation (Mookkunimala). In the open-habitat sites (Pongumoodu and Technopark), we found a more modest species richness despite higher sampling intensity. Nevertheless, there were some interesting species there too, particularly the Gaudy Baron (\u003cem\u003eEu. lubentina\u003c/em\u003e), which is protected in India under Schedule IV of the Wildlife (Protection) Act, 1972, and was not encountered in natural forests (Das, Radhamany, and Molleman 2022).\u003c/p\u003e\u003cp\u003eNotably, we do not have good natural controls to compare our urban sites with. Without human influence, the area where Thiruvananthapuram city is now situated would have been mainly lowland rainforest, a vegetation type that is almost non-existent in this region today. The nearest natural vegetation available to us was semi-deciduous forest at the foot of the Western Ghats. However, this may be similar to what would cover Mookkunimala if there had been no human influence. A year-long study in Silent Valley National Park, a mid-altitude rainforest further north in the Western Ghats, found 32 species across forest interior and edge habitats (Das, Radhamany, and Molleman 2022). Most of the species we caught were also found there, except that we observed Travancore Palmking (\u003cem\u003eA. phidippus\u003c/em\u003e), Black Rajah (\u003cem\u003eCharaxes solon\u003c/em\u003e), Tailed Palmfly (\u003cem\u003eEl. Caudata\u003c/em\u003e), Gaudy Baron (\u003cem\u003eEu. lubentina\u003c/em\u003e), Lemon Pansy (\u003cem\u003eJunonia lemonias\u003c/em\u003e (Linnaeus, 1758)), Blue Admiral (\u003cem\u003eK. canace\u003c/em\u003e), Tamil Treebrown (\u003cem\u003eLethe drypetis\u003c/em\u003e (Hewitson, 1863)), Tranvancore Evening Brown (\u003cem\u003eP. marshalli\u003c/em\u003e), and Common Nawab (\u003cem\u003eP. athamas\u003c/em\u003e), not observed in that study. Overall, we found that in the open habitat sites, we see a subset of the species found in more natural sites in the region, most of which are more typical for open habitats, but a few forest species persist as well. The lower species richness of fruit-feeding butterflies in more open habitats may reflect that this guild is generally more diverse in forests than in more open vegetation (Schmitt et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe butterfly species composition at our sites could in part be attributed to host plants. The common species at our open-habitat sites tended to be associated with either grasses (e.g. Common Evening Brown (\u003cem\u003eMe. leda\u003c/em\u003e) and Black-branded Bushbrown (\u003cem\u003eMy. mineus\u003c/em\u003e)), cultivated trees (e.g. Common Baron (\u003cem\u003eEu. aconthea\u003c/em\u003e), Tailed Palmfly (\u003cem\u003eEl. caudata\u003c/em\u003e)), or ruderal plants (e.g. Castors and Pansies (\u003cem\u003eAriadne\u003c/em\u003e and \u003cem\u003eJunonia\u003c/em\u003e species); Nitin et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The Gaudy Baron (\u003cem\u003eEu. lubentina\u003c/em\u003e) is associated with mistletoes (\u003cem\u003eDendrophthoe\u003c/em\u003e sp.). These mistletoes are often growing on mango, fig, and tamarind, trees that are common in the city as fruit or shade trees (Ravi and Thomas 2015). Thus, conserving these vegetations is an important pillar for conserving butterflies in urban regions.\u003c/p\u003e\u003cp\u003eThere are at least two takeaways from this study. Firstly, these green areas within the urban matrix do support significant butterfly diversity that can also be exploited for nature education. For example, pupils could perform butterfly trapping on school campuses or during excursions on foot. Secondly, the temporal variation in community composition shows that to compare the butterfly diversity across urban sites, they have to be sampled over extended periods. This cannot simply be avoided by sampling for a short time in the different sites simultaneously because temporal patterns may differ among sites. The rarefaction curves indicate that a suitable sample size for quantifying differences in butterfly community parameters between sites in this system would be about 500 individuals per site.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eFM performed the sampling and supervised BCA, followed by data validation, final data analysis, and writing the final version of the manuscript. BCA performed species identification and data basing, initial data analysis, and wrote the initial version of the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe thank Ullasa Kodandaramaiah for supporting the fieldwork, and Kalesh Sadasivan for advice on sampling sites, identifications, and corrections on the manuscript. For help with the fieldwork, we are grateful to Sridhar Halali, Dheeraj Halali, Jisha Vijaykumar, O. Vignesh, Bharat Parthasarathy, and Soumen Mallick. The fieldwork of FM was funded through an INSPIRE Faculty Award to Ullasa Kodandaramaiah (DST/INSPIRE/04/2013/000476), and data curation to manuscript preparation was made possible by grant 2021/43/B/NZ8/00966 from the Naradowe Centrum Nauki (National Science Centre, Poland).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData is provided within the supplementary information file\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAronson, Myla FJ, Christopher A Lepczyk, Karl L Evans, Mark A Goddard, Susannah B Lerman, J Scott MacIvor, Charles H Nilon, and Timothy Vargo. 2017. \u0026quot;Biodiversity in the city: key challenges for urban green space management.\u0026quot; \u003cem\u003eFrontiers in Ecology and the Environment\u003c/em\u003e 15 (4): 189-196.\u003c/li\u003e\n\u003cli\u003eChettry, Vishal. 2022. \u0026quot;Peri-urban area delineation and urban sprawl quantification in Thiruvananthapuram Urban Agglomeration, India, from 2001 to 2021 using geoinformatics.\u0026quot; \u003cem\u003eApplied Geomatics\u003c/em\u003e 14 (4): 639-652.\u003c/li\u003e\n\u003cli\u003eChowdhury, Shawan, Thomas Hesselberg, Monika B\u0026ouml;hm, Mir Raihanul Islam, and Upama Aich. 2017. \u0026quot;Butterfly diversity in a tropical urban habitat (Lepidoptera: Papilionoidea).\u0026quot; \u003cem\u003eOriental insects\u003c/em\u003e 51 (4): 417-430.\u003c/li\u003e\n\u003cli\u003eChowdhury, Shawan, Shihab A Shahriar, Monika B\u0026ouml;hm, Anuj Jain, Upama Aich, Myron P Zalucki, Thomas Hesselberg, Federico Morelli, Yanina Benedetti, and Anna S Persson. 2021. \u0026quot;Urban green spaces in Dhaka, Bangladesh, harbour nearly half the country\u0026rsquo;s butterfly diversity.\u0026quot; \u003cem\u003eJournal of Urban Ecology\u003c/em\u003e 7 (1): juab008.\u003c/li\u003e\n\u003cli\u003eCyriac, Susan. 2022. \u0026quot;Dichotomous classification and implications in spatial planning: A case of the Rural-Urban Continuum settlements of Kerala, India.\u0026quot; \u003cem\u003eLand use policy\u003c/em\u003e 114: 105992.\u003c/li\u003e\n\u003cli\u003eDas, Karumampoyil Sakthidas Anoop, Dhanya Radhamany, and Freerk Molleman. 2022. \u0026quot;Community structure in an isolated tropical forest biome: one year of fruit-feeding butterfly trapping in four habitats in the Western Ghats, India.\u0026quot; \u003cem\u003eDiversity\u003c/em\u003e 15 (1). https://doi.org/10.3390/d15010036.\u003c/li\u003e\n\u003cli\u003eDeVries, Philip J, Christopher A Hamm, and James A Fordyce. 2016. \u0026quot;A standardized sampling protocol for fruit-feeding butterflies (Nymphalidae).\u0026quot; In \u003cem\u003eCore Standardized Methods\u003c/em\u003e.\u003c/li\u003e\n\u003cli\u003eDylewski, Łukasz, Łukasz Maćkowiak, and Weronika Banaszak‐Cibicka. 2019. \u0026quot;Are all urban green spaces a favourable habitat for pollinator communities? 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Springer-Verlag, New York.\u003c/li\u003e\n\u003c/ol\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":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"community composition, host plant, urban biodiversity, protected species, forest","lastPublishedDoi":"10.21203/rs.3.rs-7740235/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7740235/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUrban environments can host a wide variety of wild animals and plants, potentially contributing to conservation and facilitating city-dwellers' contact with nature. In a large city in southern India, we performed long-term sampling of fruit-feeding butterflies of two sites with mainly open habitats, and more modest sampling in two closed-canopy sites. We also sampled a nearby natural forest for comparison. We caught 3625 butterflies belonging to 29 species, of which sixteen species were recorded in the open habitats, and a further five species in the closed-canopy sites in the city. The open habitats in the city featured a protected species, the Gaudy Baron (\u003cem\u003eEuthalia lubentina\u003c/em\u003e (Cramer, 1777)), and closed-canopy sites had charismatic forest species such as the Clipper (\u003cem\u003eParthenos sylvia\u003c/em\u003e (Cramer, [1775])) and Indian Nawab (\u003cem\u003ePolyura athamas\u003c/em\u003e (Drury) 1773). Community composition differed significantly among sites and sampling months. Overall, our results show that habitats within the city have fruit-feeding butterfly diversity comparable to that of natural forests, and are thus worth conserving for nature education.\u003c/p\u003e","manuscriptTitle":"Fruit-feeding butterfly diversity in a large city in southern India","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-29 19:25:12","doi":"10.21203/rs.3.rs-7740235/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-11-08T03:15:37+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-11-06T21:31:01+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-11-04T16:33:26+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-17T05:59:29+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"25283733694892227044971055865106683733","date":"2025-10-17T03:55:21+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"253267202958401390658859726248150134864","date":"2025-10-16T17:14:14+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-16T14:15:28+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"260823442280808444481066922083836516702","date":"2025-10-16T04:05:06+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"227619321853698607748933992939572900896","date":"2025-10-15T16:11:50+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"261499241485751751713639708347666652596","date":"2025-10-15T06:10:03+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"128856980180760262508460962586081555280","date":"2025-10-15T05:30:06+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-15T01:55:38+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-10-08T10:17:01+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-10-07T03:34:01+00:00","index":"","fulltext":""},{"type":"submitted","content":"Urban Ecosystems","date":"2025-09-29T09:18:51+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"ae1ae511-c14e-4fe4-b069-7c901fe2496b","owner":[],"postedDate":"October 29th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-12-22T15:59:20+00:00","versionOfRecord":{"articleIdentity":"rs-7740235","link":"https://doi.org/10.1007/s11252-025-01879-0","journal":{"identity":"urban-ecosystems","isVorOnly":false,"title":"Urban Ecosystems"},"publishedOn":"2025-12-19 15:57:06","publishedOnDateReadable":"December 19th, 2025"},"versionCreatedAt":"2025-10-29 19:25:12","video":"","vorDoi":"10.1007/s11252-025-01879-0","vorDoiUrl":"https://doi.org/10.1007/s11252-025-01879-0","workflowStages":[]},"version":"v1","identity":"rs-7740235","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7740235","identity":"rs-7740235","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}