<<

International Journal of Research ISSN NO:2236-6124

A Study on the Congregation of Adult on Non-floral Resources at Different Locations in Jalpaiguri district of ,

Panchali Sengupta1*, Narayan Ghorai2 1Department of Zoology, West Bengal State University, Berunanpukaria, Malikapur, Barasat, District-24 Parganas (North), Kolkata-700126.West Bengal, India Email id: [email protected]

2Department of Zoology, West Bengal State University, Berunanpukaria, Malikapur, Barasat, District-24 Parganas (North), Kolkata-700126.West Bengal, India email id: [email protected]

Abstract

Several instances of puddling, as reported among different herbivore , appears quite interesting. Significantly, congregation of adult butterflies at several non-floral resources (wet soil/mud, dung, bird droppings, carrion, rotten/fermenting fruits) were examined at different locations in Jalpaiguri district adjacent to the tea estates, villages and agricultural tracts. Different of papilionids and pierids congregate on wet soil patch and puddle collectively. However other species of nymphalid, lycaenid and hesperid are found to puddle individually, without associating with others on resources like excrements and carrion. Irrespective of any species newly emerged males, and aged females are found to puddle. Interestingly, each species belonging to a particular family have a specific range of puddling duration. Such specificity in puddling among species of a family could probably be associated with their need for a common nutrient.

Keywords:, congregation, hesperid, lycaenid, nymphalid, papilionid, pierid

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5877 International Journal of Research ISSN NO:2236-6124

Introduction Puddling is a widely recognised fascinating event in the life history of any herbivore arthropods except beetles targeted towards accumulation of specific micronutrient (Mollemann, 2010). Several literature illustrating the beauty of such puddling behaviour among hymenopterans (Butler, 1940; Kaspari et al., 2008; Bänziger et al., 2009), dipterans (Kaspari et al., 2008), lepidopterans (Boggs & Jackson, 1991; Smedley & Eisner, 1995; Launer et al., 1996; Boggs & Dau, 2004; Pola & García-París, 2005; Molleman, 2010), fruit flies (Hendrichs et al., 1991, 1993) and locusts (Trumper & Simpson, 1993; Shen et al., 2009) are available. Accumulation of butterflies at mud puddles, pond edges, damp soil, carrion, animal dung and bird droppings are equally interesting (Adler & Pearson, 1982; Pivnick & McNeil, 1987; Sculley & Boggs, 1996; Beck et al, 1999; Hall & Willmott, 2000). Additionally, fermenting fruits being loaded with essential carbohydrates and alcohol also help in fuelling several body processes among butterflies (Norris, 1936; Brakefield, 1994; Braby & Jones, 1995; DeVries et al., 1997; Tang et al., 2013). Among nine botanical provinces of the Indian subcontinent, the eastern are blessed with unique diversity of plants and (Das 1995). Specifically the Himalayan region of West Bengal with several national park, sanctuaries and reserve forest sustain enormous biodiversity (Allen et al., 1997; Sivakumar & Prakash, 2004; Sivakumar et al., 2006; Chettri et al., 2008; Roy et al., 2012). Although studies highlighting the diversity of butterflies from this ecoregion are available (Chowdhury & Soren, 2011; Ghorai & Sengupta 2014; Sengupta et al., 2014; Bhattacharya et al., 2016), information regarding their resource utilization still needs to be investigated. Besides depending on floral resources (i.e. nectar and pollen), butterflies engage in supplementary feeding to gain some extra benefits (Mollemann, 2010). Such a wide repertoire of feeding substrates as utilized by butterflies could probably help in expanding their already diverse gene pool in turn enriching such ecosystem. The present study has therefore been designed to identify different non-floral resources exploited by adult butterflies across different regions in Jalpaiguri district, West Bengal, India. Researchers believe that puddling intensity shows interspecific variation across different sex and age classes (Collenette, 1934; Adler, 1982; Adler & Pearson, 1982; Berger & Lederhouse, 1985; Boggs & Jackson, 1991; Launer et al., 1996; Sculley & Boggs, 1996). Accordingly, this study has also been formulated to recognize any sex and age specific preferences among butterflies engaged in foraging from such non-floral resources. The tendency of species to form associations or to feed individually at puddle sites has also been explored. The duration of feeding by each species from such substrates has been documented and any similarity in their feeding pattern has been discussed.

Study Area: The entire study was undertaken by three trained field assistants between April 2018- March 2019 at five study sites (S-I, S-II, S-III, S-IV and S-V) established at different locations adjacent to the tea estates, villages and agricultural tracts in Jalpaiguri district of West Bengal, India (Figure 1a i, 1a ii, 1a iii). S-I located in the surroundings of Satkodali village (Figure 1.b) was characterised by extensive wet patches along with dead rotten leaves and logs. S-II was situated in the nearby regions of Mathura tea garden (Figure 1.c). S-III established in Damsibad village (Figure 1.d) was found to be the most disturbed study site. Occurrence of animal dung, bird droppings, carrion and rotten/fermenting fruits were noted in this study site. S-IV located in Kohinoor Tea Estate (Figure 1.e) had extensive wet patches. S-V located in the agricultural tracts of Nurpur village (Figure 1.f) was slightly disturbed area. Importantly, carrion, animal dung, bird droppings, and rotten/fermenting fruits were also observed in this study site.

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5878 International Journal of Research ISSN NO:2236-6124

Materials & Methods: Three fixed line transects (approximately 100 m length and 8 m breadth) were established at each study site. species feeding from different non-floral resources (i.e. mud/wet soil, animal dung, bird droppings, carrion and rotten/fermenting fruits) were sampled. Butterflies which visited, settled down and remained at the resource for more than one minute were considered for this study. More importantly, butterflies were not killed or captured during the entire study to avoid disturbances caused to other species engaged in feeding. Specifically, puddling behaviour among species was confirmed, if they were observed drinking/sucking from the available resource. Such drinking behaviour was associated with the extension and uncoiling of proboscis as observed through binoculars (10 X 42). Scan sampling of butterfly species was conducted between 1000-1600 hours with 30 minutes interval. Importantly, each transect was sampled once a week. Resampling of the same transect was conducted twice, each after an interval of fifteen days. Butterfly behaviour was observed using binoculars (10X42) from a distance of three meters to reduce any disturbances. All these observations were photographed using Nikon-COOLPIX-P90 and videography was done on several occasions. Identification of species was done on the basis of sexual dimorphism (where ever possible) using published literature (Haribal Meena, 1992; Kehimkar, 2008). However in cases where sex could not be ascertained the individuals were identified only up to the species level. The mean number of butterfly species observed puddling at family and subfamily level was determined to ascertain the Resource Preference Index (RPI) at such level. Wing wear was used to determine approximately the age of the adults as wings are not repaired with age. Wing wear was rated using the 1.5-5.0 scale. Individuals were grouped into two classes: young (wing wear rating 1.5-2.0) and old (wing wear rating 4.5-5.0) for purpose of data analysis (Boggs 1987). However wing wear ratings of 2.5, 3.0, 3.5 and 4.0 were said to belong to intermediate age group. In order to recognize the extent of age and sex specific preference of butterflies, the mean number of butterflies belonging to each sex was plotted against the age of butterflies expressed as wing wear. A particular individual belonging to each species considered for focal sampling was selected on random basis. Three such observation were conducted to obtain the mean duration of puddling (expressed in seconds) of species across transects constructed at each study site. Hierarchical Clustering (using hclust function of the program-R) was used to illustrate a similarity in the duration of puddling among species across the study area.

Results: Five butterfly families representing 89 species were observed puddling from different non-floral substrates in this study (Table 1). Highest RPI was recorded for papilionidae (60.54) and (56.68) on mud/wet soil at S-I and S-II respectively. On the contrary, maximum RPI was estimated for (65.69), (27.03) and hesperidae (18.35) at S-III (only for nymphalids) and S-IV (both for lycaenids and hesperids) (Table 2). The present survey thereby demonstrated relatively high abundance of papilionids and pierids at wet soil and river bed (Table 1, 2). Preferences for amino acid enriched carrion sources by nymphalids, hesperids and lycaenids were also displayed (Table 1, 2). Importantly, excrements originating from vertebrates being loaded with nutrients also attracted butterflies (Table 1, 2).Additionally, tropical lepidopterans congregation at rotten or fermenting fruits were worth mentioning (Table 1, 2). Interesting associations of older females as compared to younger males at puddle sites were also significant (Figure 2, 3, 4, 5, 6). Several rare instances of younger females of Eurema andersoni, Catopsilia pomona, nerissa, epijarbas, Spindasis lohita and lycaenina at puddling substrates were reported. Exceptional cases of older males puddling were also documented (Enispe euthymius, thyodamas, Stibochiona nicea, consimilis, pheretima, Spindasis vulcanus, chromus, dan and salsala). However in cases where species

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5879 International Journal of Research ISSN NO:2236-6124

could not be sexually differentiated such instances of sex and age specific preferences of butterflies were not available. Additionally, the present study revealed an initial attempt to employ Hierarchical Clustering to deduce a similarity in the duration of puddling among different species of the same family. Some nymphalids with almost similar duration of feeding were observed forming clusters II and III (Figure 7). However cluster VI, VII and VIII depicted the dominance of lycaenids highlighting a close similarity in their feeding duration (Figure 7). Additionally, cluster V showed a close similarity among papilionids, while the last cluster illustrated the dominance of hesperids (Figure 7). Papilionids and pierids with almost similar feeding duration created two sub clusters in cluster IV (Figure 6). Three sub clusters each dominated by lycaenids, hesperids and nymphalids were also crafted within cluster I (Figure 7).

Discussion: Historically, the study of puddling behaviour among lepidopterans stems from their need for water in dry habitat (Adler, 1982; Larsen, 1991; Launer et al., 1996). Such an alteration in the typical foraging strategy of butterflies towards favourable water sources was probably aimed at compensating the reduced volume of nectar in flowers during drought period (Launer et al., 1996).Several micronutrients required for physiological activities (Arms et al., 1974; Boggs, 1990, 1995) are dissolved in water and obtained during puddling. Therefore puddling appears to be a means of satisfying the demand for any depleted nutrient reserve (Molleman, 2010). The mating system of a particular species, their competitive abilities, nutrient requirement along with surrounding environmental conditions are believed to determine the choice of resources being explored at puddling sites (Beck et al., 1999). More specifically, mud puddles are considered to be a popular source of sodium among lepidopterans (Pivnick & McNeil, 1987; Smedley & Eisner, 1995, 1996; Beck et al., 1999; Boggs & Dau, 2004). Association of lepidopterons with artificially induced sodium enriched baits (Smedley & Eisner, 1996; Beck et al., 1999; Boggs & Dau, 2004) and naturally created sodium containing mud puddle has been reported substantially (Boggs & Jackson, 1991; Hall & Willmott, 2000; Pola & García-París, 2005). The present study also emphasized on a similar requirement among puddling papilionid and pierid species. The acceptance of carrion and excrements (i.e. animal dung and bird droppings) by puddling species in tropical regions is also worth mentioning (Larsen, 1991; Austin & Riley, 1995; Sourakov & Emmel, 1995; Beck et al., 1999; Hall & Willmott, 2000; Hamer et al., 2006). Reports of butterflies being attracted towards such decaying matter associated with decomposing fluid are available (Reed, 1958; Payne & King, 1969; Catts & Haskell, 1990; Austin & Riley, 1995; Hall & Willmott, 2000; Hamer et al., 2006; Chin et al., 2010). Preference for amino acid enriched carrion sources by nymphalids, hesperids and lycaenids as observed in this study has also been elaborated previously (Alm et al., 1990; Erhardt & Rusterholz, 1998; O’Brien et al., 2005; Nieves-Aldrey, 2009). Several instances show carrion feeding butterflies to supplement their nitrogenous demand in an attempt to improve their flight musculature (Hill et al., 2006). A similar requirement probably explains the dominance of such species in this study. Importantly, excrements originating from vertebrates being rich in nutrients also attract butterflies (Downes, 1973; Sevastopulo, 1974; Boggs & Dau, 2004; Rima et al., 2016). Higher content of sodium in carnivore dung as compared to herbivore dung and wet soil probably explains the preference for such substrate (Boggs & Dau, 2004). Additionally, lepidopteron association with bird droppings has also been reported previously (DeVries et al., 2008, 2009; Mollenam, 2010). Such greater preference for nitrogenous sources by species as compared to nectar (Erhardt & Rusterholz, 1998; Mevi-Schutz & Erhardt, 2003) and mudpuddle (Beck et al., 1999; Boggs & Dau, 2004) are noteworthy. Significantly, butterflies on being lured by foetid substances are known to engage in

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5880 International Journal of Research ISSN NO:2236-6124

drabbing movement of proboscis (DeVries, 1987). Aggregations of tropical lepidopterans on rotten or fermenting fruits (DeVries et al., 1997; Hamer et al., 2006) are also available here. Nymphalids are known to specialize themselves by feeding from fruit juices and tree sap besides nectar (Krenn, 2008). This probably explains the dominance of such butterflies at S-III and S-V. Both sodium and nitrogenous substrates (i.e. carrion and excrements) are known to increase male mating success and female reproductive success (Dunlap-Pianka et al., 1977; Pivnick & McNeil, 1987; Boggs, 1990, 1995). The role of sodium in maintaining neuromuscular activities has also been suggested by researchers (Downes, 1973; Arms et al., 1974). Significantly, tropical humid conditions similar to this study are known to quickly deplete protein sources and in turn generate a mixture of smaller nitrogen rich molecules (Beck et al 1999). Young males foraging for sodium at puddle sites are known to transfer substantial amount of this micronutrient to their potential mates (Adler & Pearson, 1982; Pivnick & McNeil, 1987; Smedley & Eisner, 1996). Similar episodes of nuptial gift transfer to females as observed probably explain the need for puddling at sodium enriched sites by a majority of younger males. However several instances of older females feeding on such substrates were also recognised in this study. Female butterflies involved in multiple mating were known to satisfy their need for sodium in the form of nuptial gifts as received from their mates. On the contrary females engaged in single mating probably felt the need for sodium and became associated with the puddle site at younger age (Boggs & Jackson, 1991). Interesting congregation of papilionid and pierid butterflies on sodium enriched substrates (forming sub clusters within cluster-IV) were evident (Beck et al 1999). In contrast, nymphalid, lycaenid and hesperid fed individually from nitrogenous sources (i.e. animal dung, bird droppings and carrion) (creating sub clusters within cluster-I). Generation of such sub clusters (within clusters I and IV) by butterflies with similar feeding duration could probably be guided by their common feeding behaviour and similar resource requirement (Beck et al., 1999). Unfortunately lack of sufficient empirical knowledge on the feeding duration of butterflies from non floral substrates requires further investigation.

Acknowledgement: The authors would like to extend their gratitude to the Head and faculty members of the Department of Zoology, West Bengal State University, West Bengal, India for their encouragement and support during the entire study period. The guidance and help of the local people of Jalpaiguri district, West Bengal is also acknowledged.

References: 1. Adler PH (1982): Soil and puddle visiting habits of moths. J Lepid Soc. 36: 161- 173. 2. Adler PH and Pearson DL (1982): Why do male butterflies visit mud puddles? Can J Zool. 60: 322-325. 3. Allen D, Anderton J and Kazmierczak K. (1997): Report on an ornithological visit to Buxa Tiger Reserve, West Bengal, India. Forktail: 31-48. 4. Alm J, Ohnmeiss TE, Lanza J and Vriesenga L (1990): Preference of cabbage white butterflies and honeybees for nectar that contains amino acids. Oecologia 84: 53-57. 5. Arms K, Feeny P and Lederhouse RC (1974): Sodium: stimulus for puddling behaviour by tiger swallowtail butterflies, P. glaucus. Science. 185: 372-374. 6. Austin GT and Riley TJ (1995): Portable bait traps for the study of butterflies. Tropical Research. 6(1): 5-9. 7. Bänziger H, Boongird S, Sukumalanand P and Bänziger S (2009): Bees (Hymenoptera: Apidae) that drink human tears. J Kansas Entomol Soc. 135-150.

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5881 International Journal of Research ISSN NO:2236-6124

8. Baz A, Cifrián B, Martín-Vega D and Baena M (2010): Phytophagous captured in carrion-baited traps in central Spain. Bulletin of Insectology 63(1): 21- 30. 9. Beck J, Muèhlenberg E and Fiedler K (1999): Mud-puddling behaviour in tropical butterflies: in search of proteins or minerals? Oecologia 119: 140-148. 10. Berger TA and Lederhouse RC (1985): Puddling by single male and female tiger swallowtails glaucus L. (Papilionidae). J Lepid Soc. 39 (4): 339-340. 11. Bhattacharya S, Maity R, Sarkar G, Ghosh G, Mukherjee D and Mukhopadhyay, C. (2016): Socio-Environmental survey of an ecologically important forest edge hamlet in Buxa Tiger Reserve, West Bengal, India. International Letters of Natural Sciences 52: 67-83. 12. Boggs CL. (1987): Within population variation in the demography of Speyeria mormonia (Lepidoptera:Nymphalidae). Ecography. 10(3):175-184 13. Boggs CL. (1990): A general model of the role of male-donated nutrients in female insects’ reproduction. Am Nat. 136: 598-617. 14. Boggs CL. (1995): Male nuptial gifts: Phenotypic consequences and evolutionary implications. In: SR Leather, Hardie J (Eds) Reproduction, Boca Raton, CRC Press, pp 215-242. 15. Boggs CL and Jackson LA. (1991): Mud puddling by butterflies is not a simple matter. Ecol Entomol. 16: 123-127. 16. Boggs CL and Dau B. (2004): Resource specialization in puddling Lepidoptera. Environ Entomol. 33 (4): 1020-1024. 17. Borthakur U, Saini RP, Gupta SS, Jakher R, Das C,Das AK, Sarma PK, Talukdar BK and Bharali R (2013). Non-invasive genetic assessment of population status of tigers (Panthera tigris tigris) in Buxa Tiger Reserve, West Bengal, India. Int J Biodiver Conserv. 5(1): 27-32. 18. Braby MF and Jones RE. (1995): Reproductive patterns and resource allocation in tropical butterflies: influence of adult diet and seasonal phenotype on fecundity, longevity and egg size. Oikos. 72: 189-204. 19. Brakefield PM. (1994): Egg size declines with female age in the fruit-feeding tropical butterfly, Bicyclus anynana (Satyrinae). Proc Exper Appl Entomol. 5: 53- 54. 20. Butler CG. (1940): The choice of drinking water by the honeybee. J Exp Biol. 17(3): 253-261. 21. Catts EP and Haskell NH. (1990): Entomology and Death: A Procedurals Guide. Joyce’s Print Shop, Clemson, SC, pp182. 22. Champion HG and Seth SK (1968): A Revised Survey of the Forest Types of India. Government of India Press, New Delhi, pp 404 + xxvii. 23. Chettri N, Shakya B and Sharma E (2008): Biodiversity Conservation in the Kangchenjunga Landscape. Hill Side Press (P.) Ltd., Kathmandu, pp. 171. 24. Chin HC, Marwi MA, Abdullah WO, Jeffery J, Azirun MS, Dhang CC, Ishak I and Omar B. (2010): Butterflies (Lepidoptera: Nymphalidae) associated with pig carrions in . Jurnal Sains Kesihatan Malaysia. 8(1): 1-4. 25. Chowdhury S and Soren R (2011): Light attracted butterflies: a review from the Indian sub-region with an inventory from West Bengal, India. J Threat Taxa. 3(6): 1868-1871. 26. Collenette CL (1934): On the sexes of some South American moths attracted to light, human perspiration and damp sand. Entomologist. 67: 81-84. 27. Das AP. (1995): Diversity of Angiospermic Flora of Darjeeling Hills. and Biodiversity. pp 118-127. 28. DeVries PJ. (1987): The butterflies of Costa Rica and their natural history. Papilionidae, Pieridae, Nymphalidae. Princeton University Press, pp xxii+327. 29. DeVries PJ, Murray D and Lande R. (1997): Species diversity in vertical, horizontal and temporal dimensions of a fruit-feeding butterfly community in an Ecuadorian rainforest. Biol J Linn Soc. 62(3): 343-364.

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5882 International Journal of Research ISSN NO:2236-6124

30. DeVries PJ, Austin GT and Martin NH. (2008) : Diel activity and reproductive isolation in a diverse assemblage of Neotropical skippers (Lepidoptera : Hesperiidae). Biol J Linn Soc. 94: 723-736. 31. DeVries PJ, Austin GT and Martin NH. (2009): Estimating species diversity in a guild of Neotropical skippers (Lepidoptera: Hesperiidae) with artificial lures is a sampling problem. Insect Conserv Divers. 2 (2): 125-134. 32. Downes JA. (1973): Lepidoptera feeding at puddle margins, dung and carrion. J Lepid Soc. 27: 89-99. 33. Dunlap-Pianka H, Boggs CL and Gilbert LE. (1977): Ovarian dynamics in heliconiine butterflies programmed senescence versus eternal youth. Science. 197: 487-490. 34. Erhardt A and Rusterholz HP. (1998): Do peacock butterflies (Inachis io L.) detect and prefer nectar amino acids and other nitrogenous compounds? Oecologia. 117: 536-542. 35. Ghorai N and Sengupta P. (2014): Altitudinal Distribution of Papilionidae butterflies along with their larval food plants, in the East Himalayan landscape of West Bengal, India. Journal of Biosciences and Medicines. 2: 1-8 36. Hall JP and Willmott KR. (2000): Patterns of feeding behaviour in adult male riodinid butterflies and their relationship to morphology and ecology. Biol J Linn Soc. 69: 1-23. 37. Hamer KC, Hill JK, Benedick S, Mustaffa N, Chey VK and Maryati M. (2006): Diversity and ecology of carrion and fruit-feeding butterflies in Bornean rain forest. J Trop Ecol. 22(1): 25-33. 38. Haribal M. (1992): The Butterflies of Himalaya and Their Natural History. Sikkim, Sikkim Natural Conservation Foundation, Gangtok, pp. 217. 39. Hendrichs J, Katsoyannos BI, Papaj DR and Prokopy RJ. (1991): Sex- differences in movement between natural feeding and mating sites and tradeoffs between food-consumption, mating success and predator evasion in Mediterranean fruit- flies (Diptera: Tephritidae). Oecologia. 86: 223-231. 40. Hendrichs J, Lauzon CR, Cooley SS and Prokopy RJ. (1993): Contribution of natural food sources to adult longevity and fecundity of Rhagoletis pomonella (Diptera: Tephritidae). Ann Entomol Soc Am. 86: 250-264. 41. Kaspari M, Yanoviak SP and Dudley R. (2008): On the biogeography of salt limitation: a study of ant communities. Proc Natl Acad Sci USA 105: 17848- 17851. 42. Kehimkar I. (2008): The Book of Indian Butterflies. Bombay Natural History Society, Oxford University Press, New Delhi, pp xvi + 497. 43. Krenn HW (2008): Feeding behaviours of neotropical butterflies (Lepidoptera: Papilionoidea). Neue Serie 88: 295-304. 44. Larsen TB. (1991): The Butterflies of Kenya and their Natural History. Oxford University Press, Oxford, pp xxii+490 45. Launer AE, Murphy DD, Boggs CL, Baughman JF, Weiss SB and Ehrlich PR. (1996): Puddling behaviour by bay checkerspot butterflies (Euphydryas editha bayensis). J Res Lepid. 32: 45-52 46. Mevi-Schültz J and Erhardt A. (2003): Larval nutrition affects female nectar amino acid preference in the butterfly (Araschnia levana). Ecology 84: 2788- 2794. 47. Molleman F. (2010): Puddling: from natural history to understanding how it affects fitness. Entomol Exp Appl. 134(2): 107-113. 48. Nieves-Aldrey JL (2009): Bebederos de mariposas. Quercus. 276:52-55. 49. Norris MJ (1936): The feeding habits of the adult Lepidoptera (Heteroneura). Transactions of the Royal Entomological Society of London. 85: 61-90. 50. O’Brien DM, Boggs CL and Fogel ML. (2005): The amino acids used in reproduction by butterflies: a comparative study of dietary sources using

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5883 International Journal of Research ISSN NO:2236-6124

compound-specific stable isotope analysis. Physiological and Biochemical Zoology. 78(5): 819-827. 51. Payne JA and King EW. (1969): Lepidoptera associated with pig carrion. J Lepid Soc. 23(3): 191-195. 52. Pivnick KA and McNeil JN. (1987). Puddling in butterflies-sodium affects reproductive success in Thymelicus lineola. Physiol Entomol. 12(4):461-472. 53. Pola M and García-París, M. (2005). Marine puddling in Papilio polytes (Lepidoptera: Papilionidae). Fla Entomol. 88(2): 211-214. 54. R Development Core Team (2013): A language and environment for statistical computing. R Foundation for Statistical Computing Vienna, Austria 55. Reed Jr. HB. (1958): A study of dog carcass communities in Tennessee with special reference to the insects. Am Midl Nat. 59: 213-245. 56. Rima N, Meme A and Hossain MM. (2016): Puddling of butterflies in Jahangirnagar University campus and the bank of Bangshi river, Savar, Bangladesh. Jahangirnagar University J Biol Sci. 5(1): 57-70. 57. Roy US, Banerjee P and Mukhopadhyay, S. K. (2012). Study on avifaunal diversity from three different regions of North Bengal, India. Asian Journal of Conservation Biology. 1(2): 120-129. 58. Roy PB and Sah R. (2012): Economic loss analysis of crops yield due to elephant raiding: a case study of Buxa Tiger Reserve(West), West Bengal, India. J. Econ. Sustain Dev. 3(10): 83-88. 59. Sculley C and Boggs CL. (1996): Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies. Ecol Entomol. 21: 193-197. 60. Sengupta P, Banerjee KK and Ghorai N. (2014): Seasonal diversity of Butterflies and their larval food plants in the surroundings of upper Neora Valley National Park, a sub-tropical broad leaved hill forest in the East Himalayan landscape, West Bengal, India. Journal of Threatened Taxa 6(1): 5327-5342 61. Sevastopulo DG. (1974): Lepidoptera feeding at puddle-margins, dung and carrion. J Lepid Soc.: 167-168. 62. Shen K, Wang HJ, Shao L., Xiao K, Shu JP, Xu TS and Li GQ. (2009). Mud- puddling in the yellow-spined bamboo locust, Ceracris kiangsu (Oedipodidae: Orthoptera): does it detect and prefer salts or nitrogenous compounds from human urine? J Insect Physiol. 55: 78-84. 63. Sivakumar S and Prakash, V. (2004): Fauna of Protected areas-6. Water birds of Buxa Tiger Reserve, West Bengal. Zoos’ Print Journal. 19(4): 1451-1452. 64. Sivakumar S. Varghese J and Prakash V. (2006): Abundance of birds in different habitats in Buxa Tiger Reserve, West Bengal, India. Forktail. 22: 128-133. 65. Smedley SR and Eisner T.(1995): Sodium uptake by puddling in a moth. Science. 270: 1816-1818. 66. Smedley SR and Eisner T. (1996): Sodium: A male moth’s gift to its offspring. Proc Natl Acad Sci USA, 93(2): 809-813. 67. Sourakov A and Emmel TC. (1995): Bait trapping for butterflies in Kenya. Trop Lepid. 6: 1-2. 68. Tang Y, Zhou C, Chen X and Zheng H. (2013): Foraging behaviour of the dead leaf butterfly, Kallima inachus. J Insect Sci.13 (58): 1-16. 69. Trumper S and Simpson SJ. (1993): Regulation of salt intake by nymphs of Locusta migratoria. J Insect Physiol. 39: 857-864.

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5884 International Journal of Research ISSN NO:2236-6124

Table 1: List of butterfly species observed feeding from different non-floral substrates at five different study sites

Sl Species Species name S-I# S-II# S-III# S-IV# S-V# No code 1 SP1 FAMILY: m m c m - PAPILIONIDAE Subfamily: Papilioninae sarpedon (Linnaeus) 2 SP2 Graphium cloanthus m m - m - Westwood 3 SP3 Graphium antiphates m m - m - (Cramer) 4 SP4 (Esper) m m d m - 5 SP5 Graphium doson (C.& R. - - - m - Felder) 6 SP6 Graphium agamemnon m - d m - (Linnaeus)-male 7 SP7 Graphium agamemnon m - - m - (Linnaeus)-female 8 SP8 Papilio castor Westwood- m - - m - male 9 SP9 Papilio castor Westwood- - - - m - female 10 SP10 Papilio polymnestor m - - m - Cramer-male 11 SP11 Papilio polymnestor m - - m - Cramer-female 12 SP12 Papilio polytes Linnaeus- m - - m - male 13 SP13 Papilio polytes Linnaeus- m - - m - female 14 SP14 Papilio memnon m - - m - Linnaeus-male 15 SP15 Papilio memnon m - - m - Linnaeus-female 16 SP16 Papilio demoleus m - - m - Linnaeus 17 SP17 Family: Pieridae m m - m - Subfamily: Coliadinae Eurema andersoni (Moore)-male 18 SP18 Eurema andersoni - m - m - (Moore)-female 19 SP19 Catopsilia pomona m m - m - (Fabricius)-male 20 SP20 Catopsilia pomona m m - m - (Fabricius)-female

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5885 International Journal of Research ISSN NO:2236-6124

21 SP21 Subfamily:Pierinae m m - m - Ixias pyrene (Linnaeus)- male 22 SP22 Ixias pyrene (Linnaeus)- m m - m - female 23 SP23 Hebomoia glaucippe m m - m - (Linnaeus)-male 24 SP24 Hebomoia glaucippe m m - m - (Linnaeus)-female 25 SP25 Appias albina m m - m - (Boisduval)-male 26 SP26 Appias albina m m - m - (Boisduval)-female 27 SP27 Appias libythea m m - m - (Fabricius)-male 28 SP28 Appias libythea - m - m - (Fabricius)-female 29 SP29 Appias lyncida (Cramer)- m m - m - male 30 SP30 Appias lyncida (Cramer)- - m - m - female 31 SP31 m m - m - (Fabricius)-male 32 SP32 Cepora nerissa m m - m - (Fabricius)-female 33 SP33 FAMILY: - m - m - NYMPHALIDAE Subfamily: Danainae Euploea mulciber (Cramer)-male 34 SP34 Euploea mulciber - m - m - (Cramer)-female 35 SP35 Subfamily: Charaxinae - - c, d, r - c, d, bernardus dr (Fabricius)-male 36 SP36 Charaxes bernardus - - c, d - c, r (Fabricius)-female 37 SP37 Charaxes solon - - r, dr - r, dr (Fabricius)-male 38 SP38 Charaxes solon - - r - r (Fabricius)-female 39 SP39 Charaxes marmax - - c, d, r - c, d, Westwood-male r 40 SP40 Charaxes marmax - - r - c, r Westwood-female 41 SP41 Subfamily: - - d, r - r Enispe euthymius (Doubleday)-male 42 SP42 Enispe euthymius - - r - r

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5886 International Journal of Research ISSN NO:2236-6124

(Doubleday)-female 43 SP43 Subfamily: m m - m - Cethosia cyane (Drury)- male 44 SP44 Cethosia cyane (Drury)- m m - m - female 45 SP45 aoris m m - m - Doubleday-male 46 SP46 Cirrochroa aoris - m - - - Doubleday-female 47 SP47 Subfamily: Limenitinae - - r m - phemius (Doubleday)-male 48 SP48 Euthalia phemius - - - m - (Doubleday)-female 49 SP49 Subfamily: m - d m d Cyrestis thyodamas Boisduval-male 50 SP50 Cyrestis thyodamas - - m - - Boisduval-female 51 SP51 Dichorrhagia nesimachus m - d m d (Doyère)-male 52 SP52 Dichorrhagia nesimachus - - d - - (Doyère)-female 53 SP53 Stibochiona nicea (Gray)- - - d m dr male 54 SP54 Stibochiona nicea (Gray)- - - dr m - female 55 SP55 Subfamily: - - r m r (Westwood)-male 56 SP56 Euripus consimilis - - r - - (Westwood)-female 57 SP57 Subfamily: m - - m - Nymphalinae Hypolimnas misippus (Linnaeus)-male 58 SP58 Hypolimnas misippus m - - m - (Linnaeus)-female 59 SP59 FAMILY: m m dr - - LYCAENIDAE Subfamily: Curetinae Curetis thetis (Drury) 60 SP60 Subfamily: - m dr - - cippus (Fabricius)-male 61 SP61 - m - - - (Fabricius)-female 62 SP62 Deudorix epijarbas - m - m -

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5887 International Journal of Research ISSN NO:2236-6124

(Moore)-male 63 SP63 Deudorix epijarbas - m - m - (Moore)-female 64 SP64 Rapala pheretima - - - m - (Hewitson)-male 65 SP65 Rapala pheretima - - - m - (Hewitson)-female 66 SP66 Spindasis vulcanus - - dr - dr, d (Fabricius)-male 67 SP67 Spindasis vulcanus - - - - dr (Fabricius)-female 68 SP68 Spindasis lohita m - - m - (Horsfield)-male 69 SP69 Spindasis lohita - - - m - (Horsfield)-female 70 SP70 Subfamily: m m - m - Polyommatinae (C. & R. Felder)-male 71 SP71 Anthene lycaenina (C. & - - - m - R. Felder)-female 72 SP72 pactolus (C. & - - dr m dr, d R. Felder)-male 73 SP73 Nacaduba pactolus (C. & - - - - dr R. Felder)-female 74 SP74 dana (de - m - m d Nicéville)-male 75 SP75 Petrelaea dana (de - m - m d Nicéville)-female 76 SP76 strabo m m - m - (Fabricius)-male 77 SP77 Catochrysops strabo - m - m - (Fabricius)-female 78 SP78 Megisba malaya - - - - dr, d (Horsfield)-male 79 SP79 Megisba malaya - - - - dr (Horsfield)-female 80 SP80 FAMILY: m - - m - HESPERIIDAE Subfamily: Hasora chromus (Cramer)-male 81 SP81 Hasora chromus - - - m - (Cramer)-female 82 SP82 (Moore) - m dr - dr 83 SP83 Subfamily: Pyrginae m m dr - dr (Fabricius)-male 84 SP84 Pseudocoladenia dan m m - - d

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5888 International Journal of Research ISSN NO:2236-6124

(Fabricius)-female 85 SP85 Subfamily: Hesperiinae - - dr - d Telicota ancilla (Herrich- Schäffer)-male 86 SP86 Telicota ancilla (Herrich- - - - - d Schäffer)-female 87 SP87 farri (Moore) - - d - dr 88 SP88 Iambrix salsala (Moore)- - - dr - dr male 89 SP89 Iambrix salsala (Moore)- - - d - dr female # where m=mud, c=carrion, d=animal dung, dr=bird droppings, r=rotten/fermenting fruits

Table 2: Resource Preference Index (RPI) of butterfly species at family and subfamily level

Taxonomic category: S-I S-II S-III S-IV S-V Family/subfamily Papilionidae 60.04 20.85 7.05 40.57 0.00 Papilioninae 60.04 20.85 7.05 40.57 0.00 Pieridae 23.57 56.68 0.00 30.46 0.00 Coliadinae 30.50 22.47 0.00 32.21 0.00 Pierinae 69.50 77.53 0.00 67.79 0.00 Nymphalidae 8.42 7.50 65.69 11.42 54.62 Danainae 0.00 48.00 0.00 34.80 0.00 Charaxinae 0.00 0.00 50.54 0.00 60.77 Morphinae 0.00 0.00 16.99 0.00 15.90 Heliconiinae 29.88 52.00 0.00 17.58 0.00 Limenitinae 0.00 0.00 3.93 9.16 0.00 Cyrestinae 33.54 0.00 22.27 25.64 20.77 Apaturinae 0.00 0.00 6.26 3.66 2.56 Nymphalinae 36.58 0.00 0.00 9.16 0.00 Lycaenidae 5.39 11.05 15.63 15.95 27.03 Curetinae 38.09 16.30 46.62 0.00 0.00 Theclinae 20.95 42.93 41.35 62.25 46.63 Polyommatinae 40.95 40.76 11.28 37.74 53.37 Hesperiidae 2.57 3.90 11.63 1.59 18.35 Coeliadinae 34.00 46.15 20.20 100 27.48 Pyrginae 66.00 53.85 10.10 0.00 15.27 Hesperiinae 0.00 0.00 69.70 0.00 57.25

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5889 International Journal of Research ISSN NO:2236-6124

1a i 1a ii

1aiii 1b

a ii

26°16′--27°0′ N 88°4′-89°53′E 26°32'27.1"N 89°29'44.9"E

26°31'55.1"N 26°36'41.9"N 89°41'43.0"E 89°23'31.1"E

1c 1d

26°33'59.9"N 26°38'02.8"N 89°41'41.0"E 89°40'59.3"E

1e 1f

Figure 1: Map of the study area where Fig. 1ai: Map of India showing West Bengal; Fig.1aii: Map of West Bengal showing Jalpaiguri district; Fig. 1aiii: Satellite imagery of Jalpaiguri district; Fig. 1b: Satellite imagery of S-I (Satkodali village); Fig. 1c: Satellite imagery of S-II (Mathura Tea Garden); Fig.1d: Satellite imagery of S-III (Damsibad village); Fig. 1e: Satellite imagery of S-IV (Kohinoor Tea Garden); Fig. 1f: Satellite imagery of S-V (Nurpur village)

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5890 International Journal of Research ISSN NO:2236-6124

25 SP1

20

SP2

15

SP3 SP16

10 SP4 SP6 SP21 SP17 SP29 SP19 SP12 SP7 5 SP23 Number individuals of eitherof sex SP27 SP10 SP57 SP59 SP25 SP14 SP51 SP8 SP49 SP70 SP24 SP68 SP43 SP13 SP58 SP84 SP76 SP45 SP80 SP83 SP31 SP26 SP22 SP11 SP15 SP20 SP44 SP32 0 0 1 2 3 4 5 6 Age Age (expressed as wing wear) Figure 2: Relationship between number of individuals of either sex and their age (expressed as wing wear) at study site 1 (name of butterfly species are denoted by their species code as mentioned in Table 1)

18

16 SP1

14 SP2

12

SP29 SP19 uals of eitheruals of sex 10 SP21 SP23 SP25 8 SP31 SP27 6 SP26 SP30 SP17 SP20 4 SP3 SP33 SP62 SP22 SP59 SP76 SP24 SP82

Number individof SP43 SP83 SP34 2 SP63 SP28 SP4 SP45 SP70 SP46 SP18 SP60 SP75 SP77 SP44 SP32 SP61 SP84 0 SP74 0 1 2 3 4 5 6 Age Age (expressed as wing wear)

Figure 3: Relationship between number of individuals of either sex and their age (expressed as wing wear) at study site 2 (name of butterfly species are denoted by their species code as mentioned in Table 1)

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5891 International Journal of Research ISSN NO:2236-6124

10

SP35 SP39 9

8

SP41 7

SP59 6

5

4 SP1 SP53 SP36

3 SP60 SP87 SP66 SP51 SP55 SP47 SP42 2 SP37 SP82 SP40 SP49 SP50 SP72 SP38 SP88 SP89 Number individuals of eitherof sex 1 SP4 SP6 SP56 SP85 SP52 SP54 SP83

0 0 1 2 3 4 5 6 Age Age (expressed as wing wear)

Figure 4: Relationship between number of individuals of either sex and their age (expressed as wing wear) at study site 3 (name of butterfly species are denoted by their species code as mentioned in Table 1)

14

13 SP19 SP16 12

11

SP6 SP12 SP1 10 SP13 SP29 9 SP3 SP62 8 SP33 SP21 7 SP30

6 SP2 SP8 5 SP20 SP14 SP23 SP68 SP77 SP64 4 SP17 SP25 SP4 SP76 SP7

SP10 SP27 SP51 SP5 3 SP65 SP32 SP63 SP26 SP28 SP81 SP31 SP49 SP45 SP22 SP11 SP9 SP18 Number individuals of eitherof sex 2 SP24 SP47 SP75 SP70 SP34 SP80 SP58 SP44 SP43 1 SP74 SP53 SP55 SP72 SP54 SP48 SP15 SP69 SP71 SP57 0 0 1 2 3 4 5 6 Age Age (expressed as wing wear) Figure 5: Relationship between number of individuals of either sex and their age (expressed as wing wear) at study site 4 (name of butterfly species are denoted by their species code as mentioned in Table 1)

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5892 International Journal of Research ISSN NO:2236-6124

8

SP35 7

6

SP39 5 SP66

SP37 SP41 4 SP67 SP82

3 SP72 SP49 SP40 SP36

SP51 SP53

2 SP76 SP85 SP42 SP79 SP87 SP86 SP88 SP73

Number individuals of eitherof sex 1 SP83 SP74 SP55 SP75 SP84 SP38 SP89

0 0 1 2 3 4 5 6 Age Age (expressed as wing wear)

Figure 6: Relationship between number of individuals of either sex and their age (expressed as wing wear) at study site 5 (name of butterfly species are denoted by their species code as mentioned in Table 1)

Figure 7: Cluster dendrogram illustrating the duration of feeding of butterflies at different non-floral substrates (where name of butterfly species are denoted by their serial number as mentioned in Table 1)

*corresponding author Volume VIII, Issue VI, JUNE/2019 Page No:5893