Effects of weather on the behaviour of urban butterflies

Effects of weather on the behaviour of urban butterflies | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 20 November 2025 V1 Latest version Share on Effects of weather on the behaviour of urban butterflies Authors : Ravi Jambhekar 0000-0001-8587-4767 [email protected] , Saskya van Nouhuys , Jagdish Krishnaswamy , Ryan Satish , Vanshika Pal , Souradeep Dhar , Simi John , Navaneethkrishnan P S , and Jyotirmoy Behera Authors Info & Affiliations https://doi.org/10.22541/au.176366361.10806242/v1 561 views 156 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract The behavioral responses of animals to weather conditions affect their individual fitness, and thus their species ranges, and persistence of populations in the face of climate change. Extreme weather is especially challenging for species inhabiting urban environments. We observed the behavior of five common butterfly species in the tropical megacity of Bengaluru, India over one year. We followed 1011 individuals, recording the time they spent flying, feeding, resting, and basking under natural weather conditions. We found that overall feeding decreased with increasing temperature, while time spent in flight increased. Butterflies increased feeding after heavy rain. There was no overall effect of temperature anomaly (higher or lower than average temperature) or whether the observation took place during a heat wave or cold spell, suggesting that these tropical butterflies are sensitive to immediate conditions independent of the longer-term context. While these where the general trends there were also species-specific responses. For example, as temperature increased, feeding by the common four-ring, chocolate pansy, lemon pansy and tawny coster declined. In contrast, feeding by the common grass yellow increased with temperature. These findings highlight importance of species-specific behavioral plasticity in shaping butterfly resilience to ongoing climate change. This has implications for predicting future shifts in local distributions and informing urban biodiversity conservation strategies. Introduction Extreme weather events linked with climate change are known to affect biodiversity, species, populations and the behavior of individuals across the globe (Hill et al., 2021). Some species have benefited from these extreme weather events while others have been affected negatively (Pacifici et al., 2015). An important way in which weather effects species is through its impact on behaviour. Components of daily butterfly behavior such as flight, feeding, basking, resting, and ovipositing are strongly influenced by local weather and specific microclimate conditions (Kallioniemi, 2013; Kleckova et al., 2014). Depending on the local conditions butterfly individuals will engage differently in these activities, which affects their fitness. As microclimates differ due to temperature, humidity, rain or wind, so does the behavior of butterflies. In urban environments micro-climatic conditions including heat island effects may determine the ability of some species to persist (Yang et al., 2016). More extreme urban conditions are increasing with the incidence of extreme weather events globally (Ummenhofer and Meehl, 2017). Taxa with a short adult life span, such as most butterflies, are particularly vulnerable to extreme weather events because conditions over a short timescale can have a large impact (De Palma et al., 2017; McDermott Long et al., 2017). On the other hand, micro-climatic conditions may also serve as refuges from extreme climatic conditions. Hence, species responses to climate change and micro-climatic perturbations vary a lot and are difficult to predict (Hill et al., 2021). While the effects of weather on species is ultimately determined by survival or population growth rate, behind these direct measures of fitness is behavior. The changing climate and its micro-climatic impacts on behavior that is closely related to fitness is understudied, especially in the rapidly expanding and often extreme conditions of urban environments. We explore effects of microclimatic variation as a proxy for climate change on butterfly behavior in the tropical mega-city of Bengaluru, India. Insects and other invertebrates are especially vulnerable to the effects of changing climate and micro-climatic conditions because they small and ectothermic (Halsch et al., 2021). Micro-climatic change is one of the important reasons for the far reaching insect decline which has been reported (Goulson, 2019; Jarvis, 2018). However, it is difficult to study most insects because they are inconspicuous and there is little information available on their life history, the habitats they occupy, and their behaviour. Most arthropod species are even difficult to identify. Butterflies however, are relatively conspicuous, well documented and found in a diversity of habitats including forests, grasslands and even urban green areas. They are known to be affected by climate change and daily fluctuations in weather and habitat fragmentation (Hill et al., 2021; Parmesan and Singer, 2022). This makes butterflies good for behavioural and population level study. The behavioural responses of butterflies to weather and sudden changes in micro-climate are central to their adaptability to the changing climate. Butterflies are an important component of biodiversity, as herbivores, prey to birds and other insects, and as pollinators. They are also appreciated by many citizens in urban areas (McGinlay et al., 2017). In cities across the world, much of the butterfly diversity is found in gardens, parks and institutional campuses (Wang et al., 2017). Much of the literature on effects of weather on butterflies comes from temperate countries (Ashe‐Jepson et al., 2024; Warren et al., 2021). Most of these studies are reported from natural habitats and detailed studies are available only for a few model systems such as the Fritillaries (Ehrlich and Hanski, 2004; Kahilainen et al., 2018). Understanding of butterfly behavioral responses to climate change in urban areas is also lacking, though there is some evidence of butterflies behaving differently in urban than in natural habitats. A study reports that urban butterflies tend to be bold (Cormont et al., 2011), and another study found that urban butterflies tend to fly high and for a long time, which could lead to a high dispersal rate (Cormont et al., 2011; Kaiser et al., 2020). Though it has been little studied, the effects of weather on time spent flying, feeding and mating must affect population growth rate and vulnerability of species to environmental change. In this study we systematically observed the behavioral responses of five butterfly species to weather, and to extreme weather events such as heat waves, heavy rainfall events and cold spells, in an urban setting. We expected that all of the butterfly species would respond to extreme weather conditions and sudden changes in climate, and that extreme conditions would overall be negative. However, we also predicted that some species would be less adversely or even positively affected, and that some types of behaviour would respond more than others to extreme conditions. We predicted that extreme heat waves and cold spells would lead to cryptic behaviours such as resting and sitting to conserve energy. Conversely, feeding, mating and egg laying would be seen more after heavy rainfall because of increased flower availability and physiological needs of the butterflies after periods of inactivity. To address these predictions, we studied behaviour of five naturally occurring butterfly species in three locations in the large city of Bengaluru, Karnataka, India. We assessed their behaviour using focal animal sampling (Altmann, 1974) weekly throughout one year, and analysed the behavioural responses to weather, weather anomalies and long-term weather trends that occurred over this time. Methods Study site: Our study was located in Bengaluru, Karnataka. We focused on common species of butterflies found in the city that are present throughout the year. Bengaluru is in a subtropical climate with an area of 709 sq. km and a rapidly expanding population of about 20 million (Deb et al., 2020). Bangalore experiences monsoon season in June-September and a dry season period from January-March (Ravindrababu et. al 2010). This city has diverse green habitats including gardens, urban forests, water bodies, parks and campuses, and open unbuilt plots which are habitats for urban wildlife (Jambhekar et al., 2025; Swamy et al., 2019). The green parts of the city also support butterflies and many other insects pollinating and feeding on the flowering plants that grow naturally or have been planted for beautification and aesthetic purposes. Figure 1. Butterfly sampling locations in Bangalore, Karnataka, India. Maroon dots represent the 3 locations in and around the city. Green (parks, campuses and large vegetation patches) built-up areas (grey) and water (blue) are represented on the map to give an idea of the habitats of the study locations. The black line indicates the city boundary. Butterfly species: The criteria used to choose the species was that they should be common and stay close to the ground so they would be easy to follow for detailed behavioural observation. Abundances and local occurrence of each species varies but at least some individuals of each should be present year-round. We selected five species: common grass yellow ( Eurema hecabe ) Family: Pieridae, Lemon pansy ( Junonia lemonias ) Family: Nymphalidae, chocolate pansy ( Junonia iphita ) Family: Nymphalidae, tawny coaster ( Acraea terpsicore ) Family: Nymphalidae, and common four-ring ( Ypthima huebneri ) Family: Nymphalidae. The caterpillars of these butterflies feed on common small herbs in the undergrowth (Harinath et al., 2019; Rayalu et al., 2013). Lemon pansy and chocolate pansy are known to be active during the pre-monsoons and post monsoon period, and to be at low abundance during the summer months (February-May) in southern India (Harinath et al., 2019; Rayalu et al., 2013). The pansy butterflies, common grass yellow and four-ring butterflies have multiple generations per year in southern India (Harinath et al., 2019; Kim et al., 2015; Rayalu et al., 2013). All of these species have a wide distribution range (Braby et al., 2014). Study plots: The weekly survey was done in three locations, two were educational campuses and one was a large public park. The educational campuses selected were Indian Institute of Science (IISC, North Bangalore) and the Long-term Urban Ecological Observatory (LTUEO) at the Indian Institute for Human Settlements, Kengeri campus (IIHS, South West Bangalore) and the public park was Lalbagh public park (South Bangalore) (Fig. 1). We selected the three locations to be easily accessible and open throughout the year. The study plots within these three locations were open grassy unmanaged areas. We roughly demarcated areas of about 50 x 50 m for documenting butterfly behaviour. We had three 50 x 50 m plots in the IISC campus, one plot in IIHS, Kengeri campus and three plots in Lalbagh Park. Frequency of visiting the plots: We visited a plot at least once a week from May 2023-April 2024. In addition to the weekly visits, we made extra visits to these plots immediately after an extreme weather event such as heavy rainfall, or extremely hot or cold days, to monitor any differences in butterfly behaviour. Environmental variables: During each visit to a plot we recorded the local temperature and wind-speed using a Kestrel 5000 Pocket Weather Meter (NK, USA). We obtained additional rainfall and city-wide temperature weather data repositories (Funk et al., 2017; Wan et al., 2021). We have not considered season as an environmental variable in our analysis as temperature and rainfall variables are correlated with seasonality, hence we have used the direct measures of weather variables (rainfall and temperature). We calculated the cool and hot weather anomalies to see if extreme weather events strongly affect butterfly behaviours. To do this we used a time-series of daily temperature data from online weather repository, for the year 2023-24 to estimate the average annual temperature for the year. We subtracted the annual average temperature from daily temperature that we recorded in our plots to get a time-series of temperature anomalies. Other studies of temperature extremes have used a combination of methods similar to ours to estimate weather anomalies (Buckley and Huey, 2016; Ma et al., 2021). An anomaly with a positive value is when the temperature exceeded the annual average, and negative value would be cooler than average. To address the effects of weather over a longer time we used a cumulative summation function applied to the temperature anomalies in time-series. A cumulative anomalies graph over the study period indicates the duration and intensity of heat spells and cool spells (Perkins et al., 2012). We calculated the time-series of cumulative anomalies to capture the effect of longer duration heat and cold spells. As successive days of above-normal temperatures accumulate, the cumulative temperature anomaly increases toward positive values. Prolonged persistence of such high temperatures results in a period characterized by a markedly positive anomaly, indicative of a heat spell. Similarly, if cooler than average weather persists over days, we will get a similar graphical and quantitative metric defining a cool spell. The slope of the warming and cooling spells in the cumulative temperature anomalies quantify the rate of cooling or warming. The slope of cumulative temperature was estimated using the difference between successive three-day averaged temperature. For a cumulative anomaly time-series to switch signs from negative to positive or positive to negative requires a sequence of several very hot or very cold days respectively. The slope of cool or warm curve on the cumulative anomaly time-series will be proxy for the intensity of the cooling or warming episode. Behavioral observations: We recorded butterfly behaviour using focal animal sampling (Altmann, 1974; Jambhekar and Isvaran, 2016). For each observation we followed an individual for a maximum of five minutes, and a minimum of two minutes for individuals that were lost or left the plot before five minutes was completed. Anything less than two minutes was not used for the analysis. If an individual was lost before completing the five minutes the observation was stopped and observation of a new individual was started. Start and end times were recorded carefully as this was used to estimate the proportion of time spent doing different behaviours. We recorded the behaviour of an individual butterfly every thirty seconds during an observation. The behaviors recorded were (1) basking: standing with the wings open, which is associated with thermoregulation (Clench, 1966) (2) feeding: probing a flower with proboscis inserted into the flower opening (3) flying: airborne with wings open (4) mating (5) resting: sitting with wings closed. Behavioural observations were carried out between 0900-1300 hrs as this is a peak activity period for butterflies. We did not collect data if it was raining during the observation time. Each plot was walked thoroughly to find individuals of each of the target species. We tried to ensure, to the extent possible, that the same individual was not repeatedly followed during a sampling session, by following individuals in different parts of the plot. A minimum of 3-5 individuals in total were followed during a sampling occasion. Each sampling bout in a plot required a minimum of twenty minutes and maximum of forty-five minutes as we tried to follow at least one individual of each of the five species for five minutes. Analysis In order to evaluate the overall association of butterfly behaviour with weather, data from all five species were pooled together and analysed (Jambhekar and Isvaran, 2016). For each focal follow (individual observation), the proportion of time spent i. Feeding ii. Flying iii. Resting and iv. Basking was calculated as the number of instantaneous observations of the focal individual feeding, flying, resting or basking, respectively. Mating was not included because it was very rarely observed. The proportion of a follow is a measure of the relative time spent in an activity, and ranges from 0 to 1. Each of the four activities were analysed separately using a Generalised Linear Model (GLM) with binomial errors with the proportion of an observation showing that activity as the response variable, and with temperature, rainfall, windspeed, temperature anomaly and slope of the temperature trend over a three-day window to capture heat waves and cold spells of the patch as the predictors. We used species identity and location as random effects in the models. As mentioned, the slope of cumulative temperature was estimated by calculating a successive three-day average. Cumulative anomaly time-series to switch signs from negative to positive or positive to negative requires a sequence of several very hot or very cold days respectively. The magnitude of the positive or negative slope was used as a quantification of heat and cold spells indicating extreme weather events. Each focal follow was a data point in these analyses. Binomial error models were used, as the response variable was a proportion (number of behavioural scans showing a particular activity out of a given total (Crawley, 2012). We also carried out species-wise analyses using the same behavioral and environmental variables. Proportion of time spent in behavioural activities were the response variables and temperature, rainfall, windspeed, temperature anomaly, observed during the year and slope of the microclimatic variable of the patch as the predictors. We used location as a random effect in our model. Results We followed a total of 1011 individuals of five species through the year. There were 257 individual common four rings, 92 chocolate pansys, 274 lemon pansys, 279 common grass yellows, 109 tawny coasters were followed during the study duration. We spent a total of 5039 minutes following the butterflies. The temperature ranged from 18 0 -36 0 C, windspeed ranged from 0-9.4Kmph, rainfall ranged from 0-33mm on the days when observations were carried out. The anomalies fluctuated from between three degrees Celsius below four degrees Celsius above the average temperature for the region. We were also able to capture multiple heat and cold spells spanning days or weeks during the duration of our study. Generally, the weather was sub-tropical with rainfall concentrated in the months of July-September, with intermittent showers in October and November. We analysed feeding, flying, basking and resting behaviours separately (Fig. 2-3). We found that feeding increased with rainfall ( p = 0.007, df =1, X 2 = 8.4) and declined with increasing temperature ( p = 0.009, df = 1, X 2 = 28.26) (Fig. 2). The effect sizes indicated that for every millimetre increase in rainfall there was 7% increase in feeding behaviour (Estimate Odds ratio = 1.07, CI =1.02‒1.13). For every degree C rise in temperature there was a 9% decrease in feeding behaviour. Feeding behaviour was not associated with wind-speed (p = 0.09, df = 1, X 2 = 2.59), Slope in the recent weather trend ( p = 0.42, df = 1, X 2 = 0.6430), or anomaly ( p = 0.26, df = 1, X 2 = 21.58). Flying activity increased with temperature ( p = 0.0003, df = 1, X 2 = 12.89), with an 11% increase in flight with every degree rise in temperature (estimate Odds ratio = 1.11, CI = 1.05‒1.18). Other weather variables (windspeed, rainfall, anomaly and slope) did not show any statistically significant effect on flying behaviour. Figure 2. Overall effects of microclimatic weather variables on butterfly behaviour in urban areas. Resting activity decreased with increased rainfall ( p < 0.0001, df = 1, X 2 = 15.66). There was 1% decrease in resting activity with every millimetre increase of rainfall (estimate Odds ratio = 0.90, CI = 0.85-0.95). On the other hand, as anomalies in the weather increased the resting activity also increased ( p < 0.03, df = 1, X 2 = 4.62). Resting activity increased by 16% with increase in weather anomaly (estimate Odds ratio = 1.16, CI = 1.01-1.34). Other weather variables did not show any statistically significant effect on resting behaviour. Slope ( p = 0.13, df = 1, X 2 = 2.28), windspeed ( p = 0.31, df = 1, X 2 = 1.01), temperature ( p = 0.49, df = 1, X 2 = 0.47). Basking decreased as temperature increased ( p = 0.01, df = 1, X 2 = 6.27). With every degree increase in temperature there was an 8% decrease in basking activity (estimate Odds ratio = 0.92, CI = 0.86‒0.98). As anomalies in weather increased basking activity reduced ( p = 0.01, df = 1, X 2 = 5.44). With every degree increase in anomaly basking decreased by 16% (estimate Odds ratio = 0.84, CI = 0.72‒ 0.97). Other weather variables did not show any statistically significant effect on basking behaviour. Slope ( p = 0.27, df = 1, X 2 = 1.17), wind-speed ( p = 0.95, df = 1, X 2 = 0.003), rainfall ( p = 0.98, df = 1, X 2 = 0.0005). Figure 3. Effect of weather variables on butterfly behaviour Feeding, Flying, Resting and Basking. The x-axis represents the effect size. The error bars represent 95% confidence intervals. Along the y-axis is weather variables: Δ Temperature is slope of cumulative change in temperature and anomaly is the deviance from the average temperature of the year. We also analysed the behaviour of individual species. We have reported the results for two species in the text (Fig. 4). Full model predictions for each species are available in (Table A.1). Figure 4. Behaviour of Common Four-ring (I) and Lemon Pansy (II). Feeding, Flying, Resting and Basking. The x-axes represent the mean effect size. The error bars represent 95% confidence intervals. The Y-axis shows weather variables. Δ Temperature is slope of cumulative change in temperature and anomaly is the deviance from the average temperature of the year. Temperature has different effects on the behaviours of different species. As temperature increased, feeding by the common four-ring, chocolate pansy, lemon pansy and tawny coster declined (see, Fig. 4. Feeding for common four-ring, Figures A.1-3). In contrast, feeding by the common grass yellow increased with temperature (Fig. A.2). Flight by the common four-ring declined with increasing temperature, while flight by the lemon pansy increased with temperature (Fig. 4 panels I and II, Flying). Flying behaviour of chocolate pansy, grass yellow, tawny coster was not statically related to temperature (Fig. 4 panels, Table A.1, Fig. A.2-3). The common grass yellow rested less with increasing temperature, while the chocolate pansy and lemon pansy rested more as it got warmer (Fig. 4 panels). As temperature increased basking by the common four-ring, chocolate pansy, lemon pansy and tawny coster declined. Temperature did not have a significant effect on the basking and resting behaviour of common grass yellow. We were not able to analyse the basking behaviour of Tawny coster because observations of that species were sparse and basking behaviour was not observed frequently. Discussion Butterflies behave differently depending on weather (Cormont et al., 2011) and potentially also on the context in which specific weather occurs (Hayes et al., 2024). Their behavioral responses may allow them to take advantage of current conditions (Beaulieu et al., 2015), reduce the negative effects of extreme conditions (Beaulieu et al., 2015; Tsai et al., 2020), or be detrimental (Bauerfeind and Fischer, 2014; Johansson et al., 2020). We analysed the proportion of time spent feeding, flying, resting and basking under different weather conditions, and with respect to whether the current conditions are an anomaly or part of a weather trend. We found that temperature during the observation and rainfall on the day of observation affect butterfly behaviour. Other weather variables including the context in which weather occurs (anomaly and temperature trend – long-term) had effects only on some behaviours. Feeding and flying show strong responses to daily weather conditions. At higher temperatures feeding declined. With one degree rise in temperature there was a 9% decrease in feeding behaviour. This could be because they needed less energy to maintain their body temperature at higher temperatures (Pivnick and McNeil, 1985), (Niitepõld, 2019), or they avoid flying in high temperatures as high temperatures affected their physiology, such as increasing body heat during flight. In such cases, butterflies may cope by flying less or resting in cooler spots (Beaulieu et al., 2015). The above examples and explanations are from temperate countries but there might be behavioural differences depending on region and altitude. The effect of temperature on butterfly behaviour that we observed could be explained two ways 1. Decrease in feeding behaviour on hot days 2. Decrease in feeding behaviour during hot times of year. We did not analyse seasonality as an explicit variable but seasonality was captured through regular monitoring of rainfall and temperature every week. We think it could be a combination of the above effects and not a single factor acting on the butterflies. In this study higher temperatures were correlated with dry post-monsoon months during which nectaring resources are generally low which could reduce frequency of observing feeding activities. However, since we carried out our observations in semi-managed parks and garden plots where native and exotic flowering plants were present throughout the year, this low activity due to low flower availability is minimal. Further observations of feeding rates associated with nectar availability and temperature would give more clarity on what explains the behavioural differences observed. We observed that butterflies spent proportionally more time in flight at higher temperatures, likely in search of food, mates, or oviposition sites. Similar patterns have been reported elsewhere, with increased flight activity correlating with rising temperatures (Cormont et al., 2011). However most of these studies on butterfly flight are from temperate regions where higher temperatures increase metabolic rate, allowing for increased flight (Hanski, 2008; Mattila, 2015). In these temperate and Northern studies the flight and take-off temperatures are low. Such patterns are understudied in the tropics where temperature is higher (Bonebrake et al., 2010). In many tropical and subtropical regions, such as in Bangalore, higher temperatures align with dry periods when resources are scarce, possibly prompting increased flight to locate nectar or oviposition sites (Cormont et al., 2011). Additionally, butterflies might fly to seek shade or cooler microhabitats. As poikilotherms, butterflies rely on basking to raise body temperatures (Kemp and Krockenberger, 2004; Saastamoinen and Hanski, 2008). Higher ambient temperatures may reduce basking time (Akand et al., 2018; Wenda et al., 2021). Wenda et al. (2021) suggest tropical butterflies may be adapted to heat but are more vulnerable to cold, as cold spells force longer basking and less feeding, increasing exposure to low temperatures (Mazzotti et al., 2016). In our study however, even relatively low temperature had no effect on resting behaviour. Rainfall was also an important weather variable that affected the behaviour of butterflies. After rainfall and during days when there were rainy spells, butterflies spent a large proportion of their time feeding. With every millimetre increase in rainfall feeding activity increased by 7%. Correspondingly, resting was reduced after high rainfall as the butterflies were more active. In the short term this high feeding and low resting may be because they were not able to feed during the rainfall. Butterflies are also known to lay eggs when there is a new flush of leaves (Srygley et al., 2010) which in the subtropics is associated with the rainy season. Energy is required for egg production which may increase the tendency to feed during rainy days. Wind-speed did not affect feeding, flying or resting behaviours. We had expected that under very windy conditions butterflies might not fly since wind is known to reduce butterfly flight (Duplouy and Hanski, 2013; Wikström et al., 2009). We did not observe this, maybe because we did our studies in wooded areas where the windspeed was on average only 1.9Kmph (ranging from 0-9.44Kmph), on species that fly close to the ground. We also found that weather anomalies, heat waves or cold spells did not have an overall effect on butterfly behaviour, indicating that they generally responded to the immediate conditions independent of the prevailing weather. However, some individual species were affected by anomaly and cold and heat spells. A weather anomaly is defined here as a deviation in the temperature from the yearly average. Though crude, this measurement is still meaningful because we lack information on the range of temperatures that butterflies in tropical areas are able to tolerate. While most butterflies did not respond to the temperature trends there were some exceptions. For example, the common fourring fed less at high temperatures but high temperatures were during heat spells (increasing in cumulative temperature over a widow of three days) feeding behaviour increased. Such species-specific responses are intriguing but difficult to interpret without further study. During the weather experienced over this one year of monitoring, immediate temperature was important to butterflies but longer-term weather extremes were less important. This suggests a behavioral resiliency to weather extremes. Some species were more harshly affected by temperature than others, and in some cases weather anomaly and wind-speed affected their behavior. We highlight a few behaviours using Common Four-ring and Lemon pansy to compare species level patterns. These two species were seen throughout the year and in all locations. Here we chose to focus on feeding and flying as these behaviours are known to have population level consequences (Hill et al., 2021). Feeding by the common fourring, and lemon pansy declined with temperature, as it did for all the other species. Generally feeding activity of the butterflies increased with rainfall as discussed above. In contrast, feeding decreased with increasing rainfall in common grass yellow. The common grass yellow species are reliant on host plants that grow after the monsoon season (Narender, 2006). Hence, they might be adapted for laying eggs after the rains and thus feed less during the rainy season but increased feeding after the rains in order to meet the increased resource requirement required to support egg production. Generally, literature suggests that temperatures above 32 0 C are not conducive for butterfly activity (Cerrato et al., 2016). We had 12 days when the temperature during the observation was above 32 0 C, and we found that flying was very high on these days. More days with extreme temperatures could potentially lead to better understanding of sudden high/low temperatures on butterfly behaviour. Extreme weather events are associated with climate change and affecting insects globally (Cerrato et al., 2016). We found that for five common butterflies, the temperature at the time of observation and rainfall on that day affected behaviour, especially feeding and flight. Overall, the anomaly and cumulative change in temperature effects were not important, suggesting most of these butterflies have the plasticity of behaviour to withstand some weather anomalies. We observed behavioural response to weather at the time of observation that was both general over five species of butterflies, and species specific. Monitoring behaviors that affect the performance of species in the context of weather is valuable to understand the long-term consequences of global warming on butterflies in the tropics. As responses of species differ widely (Hill et al., 2021) we should be cautious about generalizations. Our study provides trends in behavioural responses in tropical butterflies which upon further exploration may have population level consequences. Applications of our study As both climate change and urbanization intensify, their interaction impact both humans and biodiversity in cities (Hill et al., 2021). Butterflies have a complex life cycle with specific requirements of host and nectar plants, and as our study shows species specific behavoral responses to weather conditions. Butterflies are conspicuous and relatively easy to identify and track. Thus, they are excellent candidates for an early warning system for environmental and ecological change in urban ecosystems as certain types of weather conditions become more or less prominent across cities (Fleishman and Murphy, 2009; Oostermeijer and Van Swaay, 1998). In addition, butterflies, especially where they occur in cultivated parks and gardens in cities, add a social-ecological context for their conservation as an indicator of broader ecological shifts that connects human wellness with biodiversity conservation. Our study showed that common butterflies in a large crowded subtropical city were mostly resilient to weather events over a year, in terms of daily fitness related behaviour. The expectation that this behavioural resilience should translate to low weather related perturbation of butterfly population sizes. However, weather can affect resource availability on seasonal time scales, and research connecting behaviour to population size is lacking. Thus this work is needed as well as investment in regular spatially explicit butterfly surveys and monitoring to help cities plan for integration of Nature-based solutions, ecosystem-based adaptation and biodiversity conservation into urban planning (Cooper et al., 2024). 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Figure 2. Overall effects of microclimatic weather variables on butterfly behaviour in urban areas. Figure 3. Effect of weather variables on butterfly behaviour Feeding, Flying, Resting and Basking. The x-axis represents the effect size. The error bars represent 95% confidence intervals. Along the y-axis is weather variables: Δ Temperature is slope of cumulative change in temperature and anomaly is the deviance from the average temperature of the year. Figure 4. Behaviour of Common Four-ring (I) and Lemon Pansy (II). Feeding, Flying, Resting and Basking. The x-axes represent the mean effect size. The error bars represent 95% confidence intervals. The Y-axis shows weather variables. Δ Temperature is slope of cumulative change in temperature and anomaly is the deviance from the average temperature of the year. Information & Authors Information Version history V1 Version 1 20 November 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords animal behaviour climate change tropical ecology urban ecology Authors Affiliations Ravi Jambhekar 0000-0001-8587-4767 [email protected] Indian Institute for Human Settlements View all articles by this author Saskya van Nouhuys Indian Institute of Science View all articles by this author Jagdish Krishnaswamy View all articles by this author Ryan Satish Indian Institute for Human Settlements View all articles by this author Vanshika Pal View all articles by this author Souradeep Dhar View all articles by this author Simi John View all articles by this author Navaneethkrishnan P S Indian Institute of Science Education and Research Kolkata View all articles by this author Jyotirmoy Behera View all articles by this author Metrics & Citations Metrics Article Usage 561 views 156 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Ravi Jambhekar, Saskya van Nouhuys, Jagdish Krishnaswamy, et al. Effects of weather on the behaviour of urban butterflies. 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