ASJ: International Journal of Wildlife and Endangered Species Conservation (IJWESC) Vol. 2 (02) 10 August, 2019, Pp. 70-87 www.academiascholarlyjournal.org/ijwesc/index_ijwesc.htm Indexed In: Directory of Research Journals Indexing - http://www.drji.org Also Available@; Internet-Archive@Vitthalrao_and_Sharad Open access

Influence of Length of Proboscis on the Feeding Rate in Butterflies (: Hesperiidae) from Mayureshwar Wildlife Sanctuary Supe Baramati (India)

1Vitthalrao Bhimasha Khyade and 2Sharad Ganpat Jagtap

1Department of Zoology, Shardabai Pawar Mahila Mahavidyalaya, Shardanagar Tal. Baramati Dist. Pune – 413115 (India). Author’s Email ✉: [email protected]

2Department of Zoology, B.E. Society Daunds Lt. K. G. Kataria College of Arts and Science, Ambedkar Chawk, Siddhtech Road, Daund, District: Pune-413801 (India). Author’s Email ✉: [email protected]

Accepted August 02, 2019

There is a significant interaction among the plant species and butterfly species. Their interaction is leading to develop favourable characters in plants as well as in the butterflies. Hesperiidae butterflies are well recognized for the significantly longer length of their proboscis of mouth parts. In the process of evolution, butterflies gained the advantage of longer proboscis. Due to presence of longer proboscis, the butterflies of family: Hesperiidae are able to visit the host plant flowers with longer tube of corolla. The present deals with study of the Hesperiidae butterflies of Mayureshwar Wildlife Sanctuary, Supe Baramati (India) with reference to size of the body; length of proboscis (mm); diameter of food canal and rate of intake of nectar from the host plant flower (nL/s). The longest proboscis among the Hesperiidae butterflies of Mayureshwar Wildlife Sanctuary, Supe Baramati (India) in present attempt belong to grass-skipper butterfly, Perichares lotus (A. Butler, 1870) (Hesperiinae Clade: 113), measuring about 63.486 (± 9.883) units. The shortest proboscis among the Hesperiidae butterflies of Mayureshwar Wildlife Sanctuary, Supe Baramati (India) in present attempt belong to dicot skipper butterfly, undulates (Hewitson, 1867) (), measuring about 12.93 (± 1.493) units. With reference to rate of intake of fluid food material (nL/S), dicot skipper butterfly, quadratum quadratum (Sepp, 1845) (Eudaminae) reported the highest readings 1282.09 (± 277.14) units.

Keywords: Grass-skipper; Typhedanus; Perichares; Bungalotis; Calpodini.

INTRODUCTION

The most ubiquitous food material favoured by behavioural traits. The mouth parts in butterflies are like birds, butterflies and bats is the nectar the specializations as adaptations for the purpose to (Nicolson, 2007). Different types of traits in the collect the nectar from flowers from the available flora organisms are supposed to the products of evolution. (Pellmyr, 2002, Muchhala and Thomson, 2009, The traits, especially the behavioural are the Johnson and Anderson, 2010, Karolyi et al., 2012, specializations for adaptations. The physiological 2013). According to Kernn, et al. (2005), the and morphological traits are reflecting on the mouthparts in butterflies are the conspicuous Vitthalrao and Sharad. 71

elongations of the appendages around pre-oral flower as well as withdrawing and re-coiling the cavity. The proboscis is nothing but, the anterior proboscis. The rate of intake of nectar per unit time extension of the part (or parts) around the mouth. determine the efficiency of feeding the butterflies. Quality in terms of length and diameter of food There is possibility of increase in the duration of channel of proboscis get reflect on the behavioural handling the flower by butterflies with longer pattern, especially for collection of nectar from host proboscis. The possible reasons for this include plant flowers. The with significantly long problems with flower manipulation, deceleration of proboscis include euglossine bees; some tabanid nectar intake or a combination of both (Heinrich, flies; nemestrinid flies and some hawk moths. In 1975; Whitham, 1977; May, 1988). these insects, the proboscis measures twice the body According to Wolf, et al. (1972), the rate of intake length (Amsel, 1938, Borrell, 2005, Borrell and of energy in the form of food material by the is Krenn, 2006, Pauw et al. 2009, Karolyi et al. 2012, directly influencing on the foraging efficiency. Intake 2014). The insects with significantly long proboscis rate during feeding influences foraging efficiency. deserve the advantage to gain access to host plant According to Hainsworth et al. (1991). Reproductive flowers with long corolla tube. Such significant long fitness and rapid feeding should therefore be proboscis is rare in butterflies. According to Paulus favoured by natural selection. Daniel et al. (1989); and Krenn (1996), the length of proboscis of the Kim et al. (2011) and Lee et al. (2014) correlate the butterflies of most of European species is medium. It nectar feeding through a tubular proboscis with the measures about two-thirds of the body length. subject to physical laws of fluid dynamics. The Neotropical Eurybia butterflies (Riodinidae) and for morphological configuration of the feeding apparatus some Neotropical skipper butterflies (Hesperiidae) and nectar viscosity modify the rate of nectar intake. have recorded significantly longer proboscis. Earlier Based on the Biophysics Principles, there are researchers (Kunte, 2007, Bauder et al. 2011, 2013, several parameters affecting the speed of fluid 2014) have recorded the proboscis length, in feeding. Therefore, principles in biophysics help to butterflies (belong to Riodinidae and Hesperiidae) understand the constraints regarding the evolution of exceeding twice the length of body. The butterflies extremely long proboscis in butterflies (Kingsolver visiting the host plant flowers with long as well as and Daniel, 1979, 1995; Lee et al., 2014). Kingsolver short corolla tube have scarcity in their significantly and Daniel (1979, 1995) explained the nectar intake longer proboscis. According to Agosta and Janzen rate by the butterflies through the physical law of (2005), the butterflies with scarcity of long proboscis Hagen-Poiseuille. Accordingly, the nectar intake rate possibly taking a competitive advantage over short- of butterflies should increase linearly with increasing proboscis butterflies. pressure difference produced by a suction pump and Role of mouth parts in the rate of feeding is well increase with the radius of the food canal to the established fact in animals (Huming birds; Bumble exponent four. bees and some flies) using nectar as food material Theoretically, it is expected to decline linearly with On the tongue of a hummingbird: Its role in the rate escalating proboscis length (Kingsolver and Daniel, and energetics of feeding (Hainsworth, 1973; 1979, 1995). According to Borrell (2007), the Hainsworth and Wolf, 1976; Inouye, 1980; Harder, butterflies must compensate for the negative 1983; Kunte, 2007; Bauder et al., 2011; Karolyi et al., influence of a long proboscis through changes in the 2013). Earlier attempts of Kunte, K. (2007) reported radius of the food canal or the size of the suction that the duration of handling the flower by the pump, or otherwise bear this cost through a butterflies with long proboscis was significantly decreased intake rate (Borrell, 2007). longer. The efficiency of harvesting the nectar from There are some studies on a few butterfly species the flower per unit time is significant and functional and other animals such as euglossine bees, constraint for evolving extraordinarily long proboscis. hummingbirds and honeyeaters (Hainsworth, 1973; According to Karoyi et al (2013), the process of Kingsolver and Daniel, 1983; May, 1985; Mitchell and handling the flower by the butterflies can be divided Paton, 1990; Molleman et al., 2005; Borrell, 2007). into the steps like, entering into the flower; taking up There are rare reports on exact measurements of the nectar and leaving the flower. Each of the step nectar intake rates combined with morphological listed deserve significant efficiency. Butterflies with quantitative data over a variety of butterfly species. longer proboscis have to manipulate the uncoiling Video recordings of foraging activity of Hesperiidae in proboscis spiral and finding an entrance into the the wild and during standardized feeding 72 Int. J. Wildlife and Endangered Species Conserv.

experiments help explain whether prolonged flower measured through the method explained by Bauder, handling times of long-proboscis butterflies result et al. (2015). According to Bauder, et al. (2013) and from decelerated nectar intake rates, prolonged Karolyi, et al (2013), measuring the exact size of flower manipulation times or both. If both of these suction pumps requires time-costly morphological behavioural aspects were independent of proboscis reconstructions. In the present attempt, it was not length, prolonged flower handling times of long- manageable for a large sample size. Therefore, the proboscis butterflies would simply result from taking present attempt preferred measuring the body size larger amounts of nectar than short proboscis as a correlate for the size of the suction pump. The butterflies. The present study presents an integrative body size is known to scale with suction pump size approach combining data obtained from behavioural (Karolyi et al., 2013). The ethanol preserved butterfly observations and morphological analyses of specimen was drained out completely and processed Hesperiidae butterflies of Mayureshwar Wildlife for pinning. The body of butterfly specimen was Sanctuary, Super Baramati India. pinned in a lateral position to a foam mat. Micrograph of body of butterfly specimen was taken through the use of a Nikon SMZ 1500 stereo-microscope (Nikon, MATERIALS AND METHODS Tokyo, Japan). After taking a micrograph of the body, the proboscis of each butterfly specimen was (A). Area of Study and Sampling of the Species: separated from the head at its base. The proboscis was then made uncoiled and fixed on a foam mat. The study was carried out through the use of pins were used for fixing the proboscis on foam Hesperiidae butterflies from Mayureshwar Wildlife mat. Sanctuary. This “Mayureshwar Wildlife Sanctuary” Micrographs of the proboscis were taken using a belong to Deccan Plateau. It is a part of Supe village Nikon SMZ 1500 stereomicroscope (Nikon, Tokyo, (Tal. Baramati Dist. Pune Maharashtra India) (Co- Japan) equipped with an Optocam-I digital camera ordinates: 180 20’ 6” N 740 22’ 15” E) (Figures 1, 2, 3 (Nikon). Micrographs of body and proboscis thus and 4). Sampling of butterflies of Family: Hesperiidae obtained were imported to Image J (U.S. National was carried out in September, October, 2018 and Institutes of Health, Bethesda USA). The length of January, February, 2019. The attempts on the body of individual butterfly and length of proboscis of morphometry and feeding by butterflies were carried respective butterfly were measured with the aid of the out through the use of three hundred thirty segmented line tool. After the length measurement, specimens. All the butterflies were belonging to the proboscis was cut off from it’s base, and the two family: Hesperiidae. Total number of species and galeae were separated from each other. one of the genera of Hesperiidae butterflies in the attempt was galea was mounted onto a microscope slide. The thirty-four and twenty-one respectively. For each food canal was kept facing upwards. It was parameter, fifty percent females and fifty percent embedded in glycerol and covered with a cover-slip. male specimens of each species were utilized. The The height of the food canal was measured using a hand net was used for the purpose to collect the Nikon Eclipse E800 light microscope (Nikon) butterflies. equipped with a Nikon Fi2-U3 digital camera (Nikon) The feeding trials were carried out for each species and the NIS Elements D software (Nikon). The width of Hesperiidae butterfly. After, the feeding trials, the of the food canal was calculated as the distance in butterflies were stored in seventy percent ethanol. micrometer between two focal planes situated on the The phylogeny explained by Warren, et al. (2009) lateral wall of the food canal and on the cuticular was followed for classification of taxa for collected spines of the dorsal linkage. The study attempt butterflies. measured the height and width of the food canal in two proboscis regions per galea, located at 10% (B) Measurements of body length; Proboscis (proximal) and 80% (distal) of the total proboscis length and diameter of food canal: length. Further, the study attempt estimated the cross-sectional area of the food canal of a proboscis For the purpose of estimation of rate of intake of in approximation to an ellipse and calculated the nectar by individual butterfly, it is necessary to know mean cross-sectional area of the proximal and distal the length of proboscis and cross-sectional area of food canal for each proboscis from respective the food canal. Both the parameters of were butterfly. Vitthalrao and Sharad. 73

https://www.researchgate.net/figure/Mayureshwar-Wildlife-Sanctuary-site_1_316084100

Figure1. Mayureshwar Wildlife Sanctuary Site.

(C) Attempts on Feeding the Butterflies: allowed for taking nectar from flowers of respective host plant. The range of average ambient air Attempts on feeding the butterflies were carried out temperatures during the attempts on feeding the in an outdoor cage (39292 m) at the Malegaon Sheti butterflies was 26oC to 30oC. The sugar solution of Farm of Agriculture Development Trust, Baramati forty percent strength was prepared. The contents of (India). The hand net collected butterflies from this solution include: Sucrose; Glucose; Fructose and Mayureshwar Wildlife Sanctuary were carried to distilled water (Alm, et al. , 1990). The artificial nectar Malegaon Sheti Farm of Agriculture Development was renewed to avoid an increase in concentration Trust, Baramati (India). Butterflies were stored in a due to evaporation. This sugar solution was placed cage until the end of the attempt on feeding. The under test conditions for half an hour to warm up to sampling session for each feeding attempt for each ambient air temperature. Pair of tweezers was used species of butterfly was minimum two hours and for pinching the wings of butterflies closed. Such maximum four hours. They were firstly made individual buttefly was placed on a feeding platform. comfortable at the place of working. Butterflies were The glass vial diameter of which was 3.64 mm was made free on fresh host plant flowers maintained in kept adjusted to the feeding platform. Glass vial was the green house at Malegaon Sheti Farm of with markings of levels of fluid. This glass vial was Agriculture Development Trust, Baramati (India). The filled with sugar solution. With the help of dissecting captured butterflies were hungry and ready to take needles, the proboscis of individual butterfly was the food material. Hungriness of butterflies is made uncoiled. Such proboscis of individual butterfly obligatory to for subsequent feeding trials. They was placed onto the sugar solution. The tip of 74 Int. J. Wildlife and Endangered Species Conserv.

Figure 2. Habitat Zonation of Mayureshwar Wildlife Sanctuary.

proboscis made inserted into the sugar solution. The Liter per second (nL/S). time of starting and ending of feeding (sucking the sugar solution) was recorded. Thus, individual (E) Flower Handling Duration Assessment: butterfly was tested once and was subsequently fixed in seventy percent alcohol. The butterflies were collected from the respective host plant flowers from the study area. The butterflies were caught from a flower shortly before they would (D) Nectar Intake Rate Assessment: start taking nectar. The butterflies were then allowed free in an outdoor cage. This cage was equipped with Through the use of software [PMB 5.0.02.11130 a freshly cut and watered inflorescence of the host (Sony Corporation)] for continuous video record of plant flower of respective butterflies (the nectar host feeding, the images were obtained. Images were plant species that butterflies had visited under natural imported to Adobe Photoshop CS4 Extended 11.0.2 conditions before being collected) (Table 2). (Adobe Systems Incorporated, San Jose, CA, USA), The video recordings were obtained separately for converted to semitransparency and overlaid. each species of butterfly for visiting host plant flower. Through the measurements of initial and final levels Analysis of video recordings of visiting the butterfly to of fluid in the vial, volume of ingested sugar solution the host plant flower were analysed through the use by individual butterfly. The reading of ingested of software PMB 5.0.02.11130 (Sony Corporation). volume of sugar solution and time required were used Assessment of behavioural patterns was carried out for calculation of rate of intake of sugar solution. The through the analysis of video record of the butterfly unit for the rate of intake of sugar solution was nano- visiting the host plant flower. Vitthalrao and Sharad. 75

Figure 3. Set-up for video-recorded feeding trials. (a) Skipper feeding from 40% sugar solution. Hungry skippers were locked into position on a stage by pinching the wings together with a pair of tweezers. The proboscis was uncoiled manually and inserted into the glass vial filled with sugar solution. (b) Measuring the ingested volume of sugar solution on video footage. The difference of fluid level from the start and the end of a feeding session was estimated in approximation to a cylinder. gv – glass vial, st – stage, tw – tweezers.

The patterns of behavioural of hesperiidae butterflies calculated using a general linear model with repeated in the attempt include: Uncoiling of proboscis; measurements (random factor: ) with the Insertion of proboscis into the corolla tube of host function lme of the package nlme (https://cran.r- plant flower; proboscis extraction and recoiling of the project.org/web/packages/nlme/nlme.pdf) (Pinheiro proboscis after finishing the feeding. The behavioural et al., 2014). Correlation between variables was patterns were assessed as manipulation time. In assessed using a Pearson correlation with the contrast, the period after successful proboscis function of the package Hmisc(Harrell, 2012). The insertion when the butterfly remained motionless was influence of proboscis length and food canal area on evaluated as the duration of suction. intake rate was calculated for a set of 21 genera using the phylogenetic comparative method as (F). Statistical Analysis of the data: implemented in the package ape (Paradis et al., 2004). This method accounts for the phylogenetic For consistency in the results, each step in the study relationship between genera as genera cannot be attempt was repeated for three times. Statistical regarded as independent from each other. package R 3.1.0 (R Development Core Team, 2011) was used for calculation of all the rests. The general linear model (with repeated measurements / random RESULTS AND DISCUSSION factor: genus) with the function lme of the package nlme (Pinheiro et al., 2014) was used for knowing the The results on the attempt are presented in Tables 1 influence of sex of butterfly on the rate of intake of and 2. The skipper (Hesperiidae) butterflies in the nectar fluid. present attempt belong to seven subfamilies: The influence of sex on rate of intake of nectar was Eudaminae; Pyrginae; Hesperiinae [Clade: 113]; 76 Int. J. Wildlife and Endangered Species Conserv.

Figure 4. Scanning Electron Micrograph of Proboscis in (Dicot Skipper Butterfly) Bungalotis quadratum quadratum (Sepp, 1845) [Family: Hesperiidae; Subfamily: Eudaminae].

Calpodini; Antoptini and . Number of species and maximum in Bungalotis quadratum quadratum of butterflies studied in the attempt was 12; 02 03; 08; (Sepp, 1845). 01; 07; and 01 belonging to subfamilies: Eudaminae; The length of the proboscis of Hesperiidae Pyrginae; Hesperiinae [Clade: 113]; Hesperiinae butterflies from Mayureshwar Wildlife Sanctuary [Calpodini]; Hesperiinae [Antoptini]; Hesperiinae Supe Baramati (India) of subfamily: Eudaminae was [Moncini] and Hesperiinae [Hesperiinae] found measured from 13.938 (± 1.829) to 39.691 (± respectively. 3.653) units. Minimum length of the proboscis in Hesperiidae butterflies from Mayureshwar Wildlife Subfamily - Eudaminae: Sanctuary Supe Baramati (India) of subfamily: Eudaminae was found in calchas (Herrich- The length of the body of Hesperiidae butterflies from Schaffer, 1869) and maximum in Bungalotis Mayureshwar Wildlife Sanctuary Supe Baramati quadratum quadratum (Sepp, 1845). (India) of subfamily: Eudaminae was found measured The food canal diameter (Square Micrometer) in from 16.297 (± 1.443) to 30.625 (± 3.864) units. Hesperiidae butterflies from Mayureshwar Wildlife Minimum body length of the body in Hesperiidae Sanctuary Supe Baramati (India) of subfamily: butterflies from Mayureshwar Wildlife Sanctuary Eudaminae was found measured from 1783.50 (± Supe Baramati (India) of subfamily: Eudaminae was 9.777) to 9155.781 (± 32.813) units. Minimum food found in Typhedanus undulates (Hewitson, 1867) canal diameter in Hesperiidae butterflies from Mayureshwar Wildlife Sanctuary Supe Baramati Pyrginae was found in in Celaenorrhinus darius (India) of subfamily: Eudaminae was found in (Evans, 1952) and maximum in Mysoria ambigua simplicius (Stoll, 1790) and maximum in (Mable and Boullet, 1908) . Bungalotis quadratum quadratum (Sepp, 1845). Rate of intake of food material (in the form of forty Subfamily – Hesperiinae [Clade – 113] : percent sugar solution) [nL/S] in Hesperiidae butterflies from Mayureshwar Wildlife Sanctuary The length of the body of Hesperiidae butterflies from Supe Baramati (India) of subfamily: Eudaminae was Mayureshwar Wildlife Sanctuary Supe Baramati found measured from 78.131 (± 19.817) to 1282.099 (India) of subfamily: Hesperiinae [Clade – 113] was (± 277.14) units. Minimum food canal diameter in found measured from 26.293 (± 1.723) to 29.968 (± Hesperiidae butterflies from Mayureshwar Wildlife 2.516) units. Minimum body length of the body in Sanctuary Supe Baramati (India) of subfamily: Hesperiidae butterflies from Mayureshwar Wildlife Eudaminae was found in in Typhedanus undulates Sanctuary Supe Baramati (India) of subfamily: (Hewitson, 1867) and maximum in Bungalotis Hesperiinae [Clade – 113] was found in quadratum quadratum (Sepp, 1845). Pyrrhopygopsis socretes orasus (H. Druce, 1876) and maximum in Perichares lotus (A. Butler, 1870). Subfamily - Pyrginae: The length of the proboscis of Hesperiidae butterflies from Mayureshwar Wildlife Sanctuary The length of the body of Hesperiidae butterflies from Supe Baramati (India) of subfamily: Hesperiinae Mayureshwar Wildlife Sanctuary Supe Baramati [Clade – 113] was found measured from 34.654 (± (India) of subfamily: Pyrginae was found measured 4.394) to 63.486 (± 9.883) units. Minimum length of from 21.253 (± 1.786) to 23.371 (± 1.963) units. the proboscis in Hesperiidae butterflies from Minimum body length of the body in Hesperiidae Mayureshwar Wildlife Sanctuary Supe Baramati butterflies from Mayureshwar Wildlife Sanctuary (India) of subfamily: Hesperiinae [Clade – 113] was Supe Baramati (India) of subfamily: Pyrginae was found in Pyrrhopygopsis socretes orasus (H. Druce, found in Celaenorrhinus darius (Evans, 1952) and 1876) and maximum in Perichares lotus ( A. Butler, maximum in Mysoria ambigua (Mable and Boullet, 1870). 1908). The food canal diameter (Square Micrometer) in The length of the proboscis of Hesperiidae Hesperiidae butterflies from Mayureshwar Wildlife butterflies from Mayureshwar Wildlife Sanctuary Sanctuary Supe Baramati (India) of subfamily: Supe Baramati (India) of subfamily: Pyrginae was Hesperiinae [Clade – 113] was found measured found measured from 15.412 (± 1.961) to 30.089 (± from 6422.47 (± 33.546) to 7756.33 (± 38.514) units. 3.829) units. Minimum length of the proboscis in Minimum food canal diameter in Hesperiidae Hesperiidae butterflies from Mayureshwar Wildlife butterflies from Mayureshwar Wildlife Sanctuary Sanctuary Supe Baramati (India) of subfamily: Supe Baramati (India) of subfamily: Hesperiinae Pyrginae was found in Mysoria ambigua (Mable and [Clade – 113] was found in Perichares adela Boullet, 1908) and maximum in Celaenorrhinus (Hewitson, 1867) and maximum in Perichares lotus ( darius (Evans, 1952). A. Butler, 1870). The food canal diameter (Square Micrometer) in Rate of intake of food material (in the form of forty Hesperiidae butterflies from Mayureshwar Wildlife percent sugar solution) [nL/S] in Hesperiidae Sanctuary Supe Baramati (India) of subfamily: butterflies from Mayureshwar Wildlife Sanctuary Pyrginae was found measured from 3468.31(± Supe Baramati (India) of subfamily: Hesperiinae 17.036) to 7323.21 (± 35.971) units. Minimum food [Clade – 113] was found measured from 558.62 (± canal diameter in Hesperiidae butterflies from 2.913) to 584.01 (± 3.145) units. Minimum food canal Mayureshwar Wildlife Sanctuary Supe Baramati diameter in Hesperiidae butterflies from (India) of subfamily: Pyrginae was found in Mayureshwar Wildlife Sanctuary Supe Baramati Celaenorrhinus darius (Evans, 1952) and maximum (India) of subfamily: Hesperiinae [Clade – 113] was in Mysoria ambigua (Mable and Boullet, 1908) . found in in Perichares lotus (A. Butler, 1870) and Rate of intake of food material (in the form of forty maximum in Pyrrhopygopsis socretes orasus (H. percent sugar solution) [nL/S] in Hesperiidae Druce, 1876). butterflies from Mayureshwar Wildlife Sanctuary Supe Baramati (India) of subfamily: Pyrginae was Subfamily – Hesperiinae [Calpodini] : found measured from 137.27 (± 10.413) to 389.83 (± 33.573) units. Minimum food canal diameter in The length of the body of Hesperiidae butterflies from Hesperiidae butterflies from Mayureshwar Wildlife Mayureshwar Wildlife Sanctuary Supe Baramati Sanctuary Supe Baramati (India) of subfamily: (India) of subfamily: Hesperiinae [Calpodini] was Vitthalrao and Sharad. 77

Table 1. The parameters affecting the rate of feeding in the skipper butterflies (Family: Hesperiidae) from Mayureshwar Wildlife Sanctuary Supe Baramati (India).

S/N Hesperiidae Butterfly Name N Size of the Length of Food Canal Rate of Intake body (mm) Proboscis Diameter (Square of Food (mm) Micrometer) Material [nL/S] 01. Eudaminae (Dicot Skipper 3 27.200 23.674 5984 486 Butterfly) (± 2.436) (± 3.417) (± 14.786) (±107.556)

Astraptes alardus latia

(Evans, 1952). 02. Eudaminae (Dicot Skipper 3 23.976 20.867 4875 527.028 Butterfly) (± 2.583) (± 3.124) (± 14.832) (±116.635)

Astraptes anaphus anetta

(Evans, 1952). 03. Eudaminae 9 17.283 16.075 3689 184.796 (± 1.323) (± 2.583) (± 13.224) (± 41.896) (Dicot Skipper Butterfly)

Autochton longipennis

(Plotz, 1882). 04. Eudaminae 6 18.938 17.619 3401 276.499 (± 1.772) (± 1.621) (± 12.189) (± 62.786) (Dicot Skipper Butterfly)

Autochton zarex

(Hubner, 1818). 05. Eudaminae 3 30.625 39.691 9155.781 1282.099 (± 3.864) (± 3.653) (± 32.813) (± 277.14) (Dicot Skipper Butterfly)

Bungalotis quadratum quadratum

(Sepp, 1845) 06. Eudaminae 9 17.566 13.938 2548.91 124.613 (± 2.216) (± 1.829) (± 11.137) (± 30.586) (Dicot Skipper Butterfly)

Cogia calchas

(Herrich-Schaffer, 1869). 07. Eudaminae 6 22.162 17.584 3777.83 243.846 (± 1.527) (± 2.022) (± 16.517) (± 61.851) (Dicot Skipper Butterfly)

Spathilepia clonius

(Cramer, 1775).

78 Int. J. Wildlife and Endangered Species Conserv.

Table 1. Continued.

08. Eudaminae 3 16.297 12.931 2778.88 78.131 (± 1.443) (± 1.493) (± 12.145) (± 19.817) (Dicot Skipper Butterfly)

Typhedanus undulates

(Hewitson, 1867). 09. Eudaminae 9 20.047 15.906 3704.55 216.71 (± 1.271) (± 1.837) (± 16.192) (± 13.741) (Dicot Skipper Butterfly)

Urbanus procne

(Plotz, 1881). 10. Eudaminae 24 17.833 15.324 1784.50 169.59 (± 1.271) (± 1.869) (± 9.777) (± 11.753) (Dicot Skipper Butterfly)

Urbanus simplicius

(Stoll, 1790). 11. Eudaminae 21 19.752 16.012 2696.25 146.97 (± 1.418) (± 1.952) (± 14.772) (± 10.185) (Dicot Skipper Butterfly)

Urbanus tanna

(Evans, 1952). 12. Eudaminae 12 19.745 16.413 2751.33 153.97 (± 1.417) (± 2.089) (± 15.073) (± 11.671) (Dicot Skipper Butterfly)

Urbanus teleus (Hubner, 1821). 13. Pyrginae 3 21.253 30.089 3468.31 137.27 (± 1.786) (± 3.829) (± 17.036) (± 10.413) (Spread-winged skippers)

Celaenorrhini

Celaenorrhinus darius

(Evans, 1952). 14. Pyrginae 12 23.371 15.412 7323.21 389.83 (± 1.963) (± 1.961) (± 35.971) (± 33.573) Pyrrhopygini

Mysoria ambigua (Mable and Boullet, 1908)

Vitthalrao and Sharad. 79

Table 1. Continued.

15. Hesperiinae 24 29.423 56.436 6422.47 567.055 (± 2.471) (± 8.786) (± 33.546) (± 1.963) Clade: 113

(Grass Skipper Butterfly)

Perichares adela

(Hewitson, 1867). 16. Hesperiinae 3 29.968 63.486 7756.33 558.62 Clade: 113 (± 2.516) (± 9.883) (± 38.514) (± 2.913)

(Grass Skipper Butterfly)

Perichares lotus

( A. Butler, 1870). 17. Hesperiinae 3 26.293 34.654 6842.15 584.01 Clade: 113 (± 1.723) (± 4.394) (± 33.974) (± 3.145)

(Grass Skipper Butterfly)

Pyrrhopygopsis socretes orasus

(H. Druce, 1876) 18. Hesperiinae 12 26.518 42.875 5597.11 538.47 (± 2.115) (± 4.421) (± 27.791) (± 2.899) Calpodini

(Grass Skipper Butterfly)

Calpodes ethlius

(Stoll, 1782) 19. Hesperiinae 6 18.536 35.461 3104.85 175.29 (± 2.201) (± 3.656) (± 320.35) (± 1.943) Calpodini

(Grass Skipper Butterfly)

Saliana esperi esperi

(Evans, 1955) 20. Hesperiinae 3 26.595 43.015 6067.43 433.17 (± 3.881) (± 4.578) (± 627.02) (± 4.762) Calpodini

(Grass Skipper Butterfly)

Saliana longirostris

(Sepp, 1840)

80 Int. J. Wildlife and Endangered Species Conserv.

Table 1. Continued.

21. Hesperiinae 9 23.472 47.548 5235.31 213.78 (± 1.664) (± 5.742) (± 696.09) (± 53.517) Calpodini

(Grass Skipper Butterfly)

Saliana salius (Cramer, 1775) 22. Hesperiinae 3 31.819 55.683 9147.92 802.99 (± 2.978) (± 7.543) (± 618.15) (± 211.01) Calpodini

(Grass Skipper Butterfly)

Saliana severus

(Mable, 1895) 23. Hesperiinae 18 21.861 41.606 4265.37 176.09 (± 1.516) (± 5.636) (± 288.22) (± 47.274) Calpodini

(Grass Skipper Butterfly)

Saliana triangularius

(Kave, 1914) 24. Hesperiinae 3 26.192 45.836 8231.33 351.57 (± 1.564) (± 2.774) (± 1183.95) (± 35.916) Calpodini

(Grass Skipper Butterfly)

Talides hispa

(Evans, 1955) 25. Hesperiinae 3 26.695 41.529 7869.09 478.53 (± 2.645) (± 2.513) (± 848.88) (± 49.889) Calpodini

(Grass Skipper Butterfly)

Thracides phidon

(Cramer, 1779) 26. Hesperiinae 3 13.881 14.286 1820.71 150.96 (± 1.806) (± 1.587) (± 196.41) (± 17.366) Anthoptini

(Grass Skipper Butterfly)

Corticea lysias lysias

(Plotz, 1883)

Vitthalrao and Sharad. 81

Table 1. Continued.

27. Hesperiinae 6 14.786 17.551 1584.90 69.139 (± 1.824) (± 1.948) (± 170.97) (± 17.348) Moncini

Cymaenes alumna (A. Butler, 1877).

28. Hesperiinae 24 15.111 21.971 2012.36 76.287 (± 1.734) (± 2.076) (± 118.54) (± 22.298) Moncini

Morys geisa

(Moschler, 1879). 29. Hesperiinae 6 15.011 19.746 2327.21 118.87 (± 1.924) (± 1.528) (± 169.35) (± 26.058) Moncini

Morys micythus

(Godman, 1990).

30. Hesperiinae 30 13.902 21.783 1887.44 71.773 (± 2.528) (± 1.685) (± 328.37) (± 19.667) Moncini

Papies phaeomelas (Hubner, 1831).

31. Hesperiinae 3 17.251 20.398 1650.72 124.91 (± 2.042) (± 1.651) (± 323.08) (± 33.227) Moncini

Papies phainis

(Godman, 1900).

32. Hesperiinae 36 22.792 32.110 2769.01 90.663 (± 1.982) (± 1.762) (± 416.46) (± 33.823) Moncini

Papies subcostulata

(Herrich-Schaffer, 1870). 33. Hesperiinae 6 15.992 16.369 1371.21 36.515 (± 2.478) (± 3.221) (± 54.583) (± 6.297) Moncini

Vehilus stictomenes iludens

(Mable, 1891).

82 Int. J. Wildlife and Endangered Species Conserv.

Table 1. Continued.

34. Hesperiinae 15 20.651 19.014 3038.46 151.11 (± 2.604) (± 2.113) (± 1.723) (± 33.991)

Pompeius pompeius

(Latrelle, 1824).

- Each Figure is the mean of three replications. - Figures with ± sign in parentheses are the standard deviations.

Table 2. Hesperiidae Butterflies used for “Flower Handling Duration Assessment” in the attempt.

S/N Host Plant: Stachytarpheta frantzii Host Plant: Calathea crotalifera 1. Autochton longipennis (N=12). Damas clavus (N=18). 2. Urbanus teleus (N=9). Saliana triangularis (N=15). 3. Morys geisa (N=6). -

N: Number of Hesperiidae Butterflies Utilized.

found measured from 18.536 (± 2.201) to 31.819 (± (Evans, 1955) and maximum in Saliana severus 2.978) units. Minimum body length of the body in (Mable, 1895). Hesperiidae butterflies from Mayureshwar Wildlife Rate of intake of food material (in the form of forty Sanctuary Supe Baramati (India) of subfamily: percent sugar solution) [nL/S] in Hesperiidae Hesperiinae [Clade – 113] was found in Saliana butterflies from Mayureshwar Wildlife Sanctuary esperi esperi (Evans, 1955) and maximum in Supe Baramati (India) of subfamily: Hesperiinae Saliana severus (Mable, 1895). [Calpodini] was found measured from 175.29 (± The length of the proboscis of Hesperiidae 1.943) to 802.99 (± 211.01) units. Minimum food butterflies from Mayureshwar Wildlife Sanctuary canal diameter in Hesperiidae butterflies from Supe Baramati (India) of subfamily: Hesperiinae Mayureshwar Wildlife Sanctuary Supe Baramati [Calpodini] was found measured from 35.461 (± (India) of subfamily: Hesperiinae [Calpodini] was 3.656) to 55.683 (± 7.543) units. Minimum length of found in Saliana esperi esperi (Evans, 1955) and the proboscis in Hesperiidae butterflies from maximum in Saliana severus (Mable, 1895). Mayureshwar Wildlife Sanctuary Supe Baramati (India) of subfamily: Hesperiinae [Calpodini] was Subfamily – Hesperiinae [Anthoptini] : found in Saliana esperi esperi (Evans, 1955) and maximum in Saliana severus (Mable, 1895). The present attempt is reporting Corticea lysias The food canal diameter (Square Micrometer) in lysias (Plotz, 1883) as only one member of subfamily: Hesperiidae butterflies from Mayureshwar Wildlife Hesperiinae [Anthoptini]. The readings 3.881 (± Sanctuary Supe Baramati (India) of subfamily: 1.806) units; 14.286 (± 1.587) units; 1820.71(± Hesperiinae [Calpodini] was found measured from 196.41) units and 150.96 (± 17.366) units belongs to 3104.85 (± 320.35) to 9147.92 (± 618.15) units. the length of the body; the length of the proboscis; Minimum food canal diameter in Hesperiidae the food canal diameter (Square Micrometer) and butterflies from Mayureshwar Wildlife Sanctuary rate of intake of food material (in the form of forty Supe Baramati (India) of subfamily: Hesperiinae percent sugar solution) [nL/S] respectively in [Calpodini] was found in Saliana esperi esperi Corticea lysias lysias (Plotz, 1883) subfamily: Vitthalrao and Sharad. 83

Hesperiinae [Anthoptini]. (Latrelle, 1824) as only one member of subfamily: Hesperiinae [Hesperiini]. The readings 20.651 (± Subfamily – Hesperiinae [Moncini]: 2.604) units; 19.014 (± 2.113) units; 3038.46 (± 196.41) units and 151.11 (± 33.991) units belongs to The length of the body of Hesperiidae butterflies the length of the body; the length of the proboscis; from Mayureshwar Wildlife Sanctuary Supe Baramati the food canal diameter (Square Micrometer) and (India) of subfamily: Hesperiinae [Moncini] was rate of intake of food material (in the form of forty found measured from 13.902 (± 2.528) to 22.792 (± percent sugar solution) [nL/S] respectively in 1.982) units. Minimum body length of the body in Pompeius pompeius (Latrelle, 1824) subfamily: Hesperiidae butterflies from Mayureshwar Wildlife Hesperiinae [Hesperiini]. Sanctuary Supe Baramati (India) of subfamily: Result analysis of present attempt reporting higher Hesperiinae [Moncini] was found in Papies rate of intake of fluid solution as a food material by phaeomelas (Hubner, 1831) and maximum in Papies the Hesperiidae butterflies with longer length of subcostulata (Herrich-Schaffer, 1870).. proboscis in comparison with the Hesperiidae The length of the proboscis of Hesperiidae butterflies with shorter length of proboscis. The rate butterflies from Mayureshwar Wildlife Sanctuary of intake of fluid solution as a food material by the Supe Baramati (India) of subfamily: Hesperiinae Hesperiidae butterflies is also influenced by the area [Moncini] was found measured from 16.369 (± of food canal, the proboscis. Both, the length of 3.221) to 32.110 (± 1.762) units. Minimum length of proboscis and area of food canal in Hesperiidae the proboscis in Hesperiidae butterflies from butterflies serve as crucial parameters for feeding Mayureshwar Wildlife Sanctuary Supe Baramati rate. The long proboscis in the Hesperiidae butterflies (India) of subfamily: Hesperiinae [Moncini] was of Mayureshwar Wildlife Sanctuary Supe Baramati found in Vehilus stictomenes iludens (Mable, 1891) (India) is closely interlinked with ecological and maximum in Papies subcostulata (Herrich- adaptations that enable an efficient nectar uptake. Schaffer, 1870). Morphological traits (body size); Anatomical traits The food canal diameter (Square Micrometer) in (musculature) and physiological traits are also Hesperiidae butterflies from Mayureshwar Wildlife important. These traits are serving for development Sanctuary Supe Baramati (India) of subfamily: of efficiency of butterflies, which probably enables the Hesperiinae [Moncini] was found measured from development of a large suction pump to overcome 1371.21 (± 54.583) to 2769.01 (± 416.46) units. nectar flow resistance. Minimum food canal diameter in Hesperiidae The evidence for morphological adaptations that butterflies from Mayureshwar Wildlife Sanctuary allow for efficient nectar intake comes from long- Supe Baramati (India) of subfamily: Hesperiinae proboscis of Eurybia butterflies (Bauder et al., 2013). [Moncini] was found in Vehilus stictomenes iludens These butterflies possess larger dilator muscles of (Mable, 1891) and maximum in Papies subcostulata the suction pump in relation to the head capsule (Herrich-Schaffer, 1870). volume compared to related short-proboscis Rate of intake of food material (in the form of forty metalmark species (Bauder et al., 2013). These percent sugar solution) [nL/S] in Hesperiidae muscles account for the occurrence of a pressure butterflies from Mayureshwar Wildlife Sanctuary drop to transport fluid into the gut (Eberhard and Supe Baramati (India) of subfamily: Hesperiinae Krenn, 2005). [Moncini] was found measured from 36.515 (± In addition, Eurybia butterflies were also shown to 6.297) to 124.91 (± 33.227) units. Minimum food have relatively large food canals (Bauder et al., canal diameter in Hesperiidae butterflies from 2013). Behavioural analyses of skippers during Mayureshwar Wildlife Sanctuary Supe Baramati flower visitation confirmed the results of Kunte (2007) (India) of subfamily: Hesperiinae [Moncini] was by showing that, the butterflies with long proboscis found in in Vehilus stictomenes iludens (Mable, 1891) require a longer time for a flower visit. and maximum in Papies subcostulata (Herrich- Further, long-proboscis Hesperiidae species of Schaffer, 1870). butterflies spent more time for drinking nectar from a flower. These findings indicate that Hesperiidae Subfamily – Hesperiinae [Hesperiini] : butterflies with longer proboscis take higher nectar volumes from the flower with deep corolla tube in The present attempt is reporting Pompeius pompeius Calathea crotalifera than butterflies with shorter 84 Int. J. Wildlife and Endangered Species Conserv.

proboscides from the flowers of Stachytarpheta larval food (Schemske and Horvitz, 1984; DeVries, frantzii. As the corolla tube of Calathea crotalifera is 1997). Some larval instars of butterflies of family: deeper than that of Stachytarpheta frantzii, the Hesperiidae feed on the leaves of Calathea species Hesperiidae butterflies that visit Calathea flowers (Schemske and Horvitz, 1984). most likely ingest higher amounts of nectar. The The larvae of the long-proboscis Hesperiidae flowers with deep corolla tube are known to secrete species are reported to feed on several more nectar than the flowers with shorter corolla tube monocotyledons (Janzen and Hallwachs, 2009) that (Harder, 1985; Harder and Cruzan, 1990). Presence occur in the understory of the forest (Weber et al., of long proboscis can be regarded as an advantage 2001), including Marantaceae (Calathea sp., because it enables butterflies to gain access to highly Marantasp., Thaliasp.), Costaceae (Costussp.), rewarding flowers. However, the flower manipulation Heliconiaceae (Heliconiasp.) and Zingiberaceae times of butterflies increased with proboscis length. (Renealmia sp.) (Janzen and Hallwachs, 2009). The Long manipulation times can lower the energy intake convergent evolution of long proboscides in rate by decreasing the proportion of foraging time Neotropical deep-forest butterfly species would devoted to actually imbibing nectar (Heinrich, 1983; provide these butterflies exclusive access to deep- May, 1985). Therefore, longer manipulation times tubed flowers, which occur in their microhabitat and could constitute functional costs of long proboscis. cannot be exploited by the vast majority of other The problem may be due to a poor supply of butterflies with shorter proboscides. Gaining the mechano- or chemosensory information, as other access to large amounts of nectar concealed inside long-proboscis butterflies (Riodinidae) are endowed long corolla tubed flowers through a long proboscis with significantly fewer sensilla on their proboscis could also serve to fulfil high energy demands of than related short proboscis species (Bauder et al., some butterflies of family: Hesperiidae. 2011, 2013). Alternatively, longer manipulation times of long-proboscis Hesperiidae butterflies could also be due to differences in flower morphology: long- CONCLUSION proboscis Hesperiidae butterflies preferred the flower with long corolla tube of Calathea crotalifera. The The allometric scaling of proboscis length with body Hesperiidae butterflies with shorter proboscis visited size virtually comes at no costs in terms of decreased flowers of Stachytarpheta frantzii with shorter corolla nectar intake rates. The highest nectar intake rate tubes. (1467 nL/s) is achieved by the only long-proboscis The bumble bees require more time to learn species featuring an almost isometric scaling complex flower designs, such as long corolla tubes relationship of proboscis length and body size. The with concealed nectar, than simple designs (Laverty, dicot skipper butterfly, Bungalotis quadratum 1994). The individual experience gained by quadratum (Sepp, 1845) (Eudaminae N=3) in the butterflies in successive attempts to forage on a present attempt reported the highest rate of intake of flower can shorten flower manipulation time (Lewis, sugar fluid. This exceptional result reported by dicot 1986; Kandori and Ohsaki, 1996; Goulson et al., skipper butterfly, Bungalotis quadratum quadratum 1997). It clearly means learning the floral morphology (Sepp, 1845) is definitely due to efficiency acquired by the butterflies could serve as an adaptive strategy through longer proboscis, body size and food canal for increasing the efficiency of nectar collection diameter. The suggests that the scaling relationship (Kandori and Ohsaki, 1996). Further, long-proboscis of body size and proboscis length may influence the butterflies could compensate for long manipulation nectar intake rate of a butterfly, which can be times by visiting fewer nectar-rich flowers instead of maximized in a large specimen featuring an isometric many flowers with less volume of the nectar. scaling relationship. These findings point to the In some Hesperiidae butterflies, for example the importance of inter-specific variation in scaling calpodines, long proboscis may have proven to be relationships and its effect on the foraging efficiency useful adaptations to their particular habitat: the deep of a butterfly. forests. These butterflies are known to live in shady, forested habitats (Warren et al., 2009). Furthermore, the long-proboscis Eurybia species of butterflies are ACKNOWLEDGEMENT also known to use the flowers of these plants not only as a nectar source of the adult butterflies, but also as Authors extend their sincere thanks to Administrative Vitthalrao and Sharad. 85

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