From the present study it was revealed that, there are nineteen different types of butterflies dependent on Lantana camara in the study area, within these months. The dominance of Papilionidae butterflies were seen from the results. Important to note that, Nymphalidae butterflies share a very good number in the community, followed by Papilionidae, Pieridae and Lycaenidae butterflies. All these butterflies show a greater preference on yellow coloured flowers, followed by white and pink flowers on a single floret. So many potential nectar plants were present, but they choose Lantana mostly. The biochemical aspects of the nectar sugar concentration and the nectar quality analysis of the Lantana plant is a thirst area for further research purpose. The present work also shows that, the proboscis lengths of the butterfly vary within and between same and different families.

1. Foraging Patterns of Some Common
Butterflies on Lantana camara - An
Exotic, Invasive Species.
DISSERTATION WORK SUBMITTED FOR THE PARTIAL
FULFILMENT FOR THE DEGREE OF MASTER OF SCIENCE IN
ZOOLOGY
BY
DEBOJYOTY GHOSH
REGISTRATION NO. 2014015667
OPTIONAL ELECTIVE: FIELD BIOLOGY
DEPARTMENT OF ZOOLOGY
WEST BENGAL STATE UNIVERSITY

2. DEDICATED WITH UTMOST LOVE & RESPECT TO
SRI GOPAL CHANDRA GUCHAIT, FROM WHOM I HAVE
LEARNED HOW TO LOVE BIOLOGY.

3. CONTENTS
INTRODUCTION …………………………………….
OBJECTIVE……………………………………………
METHODOLOGY…………………………………….
RESULT ………………………………………………..
DISCUSSION…………………………………………..
CONCLUSION………………………………………...
ACKNOWLEDGEMENT…………………………….
REFERENCES ………………………………………...

4. TO WHOM IT MAY CONCERN
This is to certify that Mr. Debojyoty Ghosh (Registration No. 2014015667 of
2014), a student of MSC, in the academic year session 2017-2019, has worked on
the project entitled “ FORAGING PATTERN OF SOME COMMON
BUTTERFLIES ON LANTANA CAMARA, AN EXOTIC, INVASIVE
SPECIES” for the partial fulfillment of Master of Science in Zoology ,under my
supervision. To the best of my knowledge this work has not been submitted to any
other universities for the award of any other degree.
Date : ………….. ……………………………………
Prof. Narayan Ghorai
Head, Department of Zoology,
West Bengal State University.

5. DECLARATION
I, Debojyoty Ghosh hereby declare that the dissertation entitled “FORAGING
PATTERN OF SOME COMMON BUTTERFLIES ON LANTANA
CAMARA, AN EXOTIC, INVASIVE SPECIES” submitted to the
Department of Zoology, West Bengal State University, for the partial
fulfillment of the degree of Master of Science in Zoology is my original work
and the dissertation has not been submitted for the award of any degree, a
diploma, to any other University or Institution.
……………………………………
Signature of the student

6. INTRODUCTION
Evolutionary interaction is found among different organisms of both flora and fauna,
generally described as co-evolution in biological terms. Various researchers worked on host-
plant and insect interactions. Both of the plants and animals evolved together, so there are
many complex relations exist between this. Some relationships are beneficial to both parties,
some are not. Some important plant-animal interactions are explored by researchers; viz.
plant pollinator, plant-herbivore, plant-disperser and some mutualism. Among all these,
plant-pollinator interaction is well studied in various aspects (Dafni, 1994). There are several
pollinators viz. insects, birds, bats found in natural system. Among all these insects,
butterflies play a major role in pollination. Nowadays, the age-old relationship between plants
and pollinators is threatened by anthropogenic causes, like habitat degradation, uses of
fertilizers and pesticides. International Union for Conservation of Nature and Natural
Resources (IUCN) defines Alien Invasive Species as an alien species which becomes
established in natural or semi- natural ecosystems or, an agent of change, and threatens native
biodiversity. These invasive are widely distributed in all kinds of ecosystems throughout the
world. The threat to biodiversity due to invasive alien species is considered second only to
that of habitat destruction. Invasive species cause loss of biodiversity including species
extinctions, and changes in hydrology and ecosystem function. Differences between native
and exotic plant species in their requirements and modes of resource acquisition and
consumption may cause a change in soil structure, its profile, decomposition, nutrient content
of soil, moisture availability etc. (Raghubanshi et al. 2005).
The word Lantana camara derives from Latin ‘lento’ which means to bend (Ghisalberti,
2000). It is the genus of verbenaceae family with 600 varieties existing worldwide. Lantana
camara, a native species of South and Central America and the Caribbean islands (Baars,
2002). Lantana camara, also known as wild sage, is a thorny multi-stemmed, deciduous
shrub with an average height of 2m (6ft). Stems are square in outline, covered with bristly
hairs when green, often armed or with scattered small prickles. Lantana camara possesses a
strong root system. The leaves of Lantana camara have a strong aroma. Its flowers are small,
multi-colored, in stalked, dense in flat-topped clusters with a corolla having narrow tube with
four short spreading lobes. Their flowers undergo color change subsequent to anthesis. These
flowers occurs in cluster which includes white-pink-lavender or yellow-orange-red mix. The
yellow coloration of the flower provides visual cue to pollinators and change in color is
initiated on the act of pollination. Lantana camara’s widespread and diverse distribution is a
reflection of its wide ecological tolerances. The species also thrive well in disturbed areas
which includes roadside, railway tracks and canals (Sharma et al., 2005). Lantana camara
have many negative impacts including potential to disrupt succession cycle, displacing native
biota resulting in decreased biodiversity (Murali and Setty, 2001). Its infestations alter the
structural and floral composition of native communities (Sharma and Raghubanshi, 2010). As
the density of Lantana camara in forest increases, allelopathic interactions increase and hence
there is decline in species richness (Day et al., 2003). The weed poses a serious problem to
flora and fauna because of its toxic substance (Lantadene A) and it contains certain
allelopathic compounds (Jain et al. 1989).
Invasive plants take common strategies to propagate, one of them are pollination; producing
very good quality of nectar or in quantity these plants attract insects. Among all insects,

7. butterflies are well known pollinators. Butterflies prefer vibrant colours, pattern of
inflorescence, tube-like corolla structure and fragrance from flowers or florets (Sengupta and
Ghorai, 2013). Butterfly posseses different innate colours, which vary between different
genera within a single family, between species in a single genus and even within or between
opposite sex in a single species (Sengupta and Ghorai, 2013).Visits of butterflies are more
frequent to flowers with tubular corollas than to non-tubular ones, to flowers of herbs and
shrubs rather than trees, to flowers which are coloured red, yellow, blue and purple than those
coloured white and pink (Tiple et al., 2006).
In India, Lantana camara was introduced as an ornamental plant, and now it has become a
widespread invasive alien plant species. In comparison with many other shrubs and
associated plants in different study sites, the relative load of butterfly was significantly higher
for L. camara. At least 25 butterfly species were found to be associated with L. camara in
varying numbers depending on the urban and rural gradient. The relative abundance of the
butterflies remained proportional to the density of flowers of L. camara irrespective of the
sites (Mukherjee et al., 2015).
An allometric relationship was found in between body size and the proboscis length of some
butterflies and flower handling time per inflorescence was well studied (Kunte, 2007).
Species with greater relative proboscis length had up to 3-times longer flower handling time
(Kunte, 2007). Important to note that, raw proboscis length have no relationship with
foraging efficiency (Kunte, 2007). Studies have considered bio-mechanical control on
foraging efficiency based on proboscis length specific studies (Borrell, 2006, 2007).
• Rational of the Study:
Being a noxious weed it keeps a good quality of faunal diversity and thereby sustains an
ecosystem service. The present study is focused on some common butterflies and their
foraging pattern in different months with their varying proboscis length with floral
morphometry on a single invasive species Lantana camara.
OBJECTIVES
1. Establishing a dominance hierarchy of some common butterflies on Lantana camara.
2. Variation between butterfly encounters in different times and in different months on
Lantana camara.
3. Flower preference of butterflies on Lantana camara.
4. Find a relationship between butterfly proboscis length and flower morphometry.

8. METHODOLOGY
The entire study was done for a period of four months, from February 2019 to May 2019.
Observations were recorded on regular basis using ad libitum and continuous scan sampling
technique (Altmann, 1974). 22 patches of Lantana camara were found in that study plot,
associated with some vegetation. The encounters of butterflies were observed diurnally in
between 9 AM to 1 PM. 1 hour scan sampling was sub-divided into four halves, each having
15 minutes. Thereby, 4 hour of intense observation was carried out in daily basis. A mobile
device (ASUS Zenfone MAX) was there to maintain the time records for scan sampling and
field photography. Butterflies were identified by using field guide books (Kunte, 2000 and
Smetacek, 2017, NCBS database). By using GPS TOOL software all environmental
parameters (viz. temperature, rainfall, and lux) and coordinates were taken.
The study site was located nearby at Khardaha Railway station, beside Electrosteel Casting
Ltd. (coordinate: 22.43.10N, 88.22.35 E).
FIGURE 1. Location map and the GPS map of the study site.
FIGURE 2: Glimps of raw data sheet.
Butterflies were collected from the study site for proboscis morphometry by using insect nets
(mesh size 2 mm.). Flowers were collected from the study site by random hand picking,from

9. 22 different patches. Proboscis morphometrical studies and corolla structure of flowers were
studied in laboratory condition, by using microscopes (device- Leica) and centimeter scales.
GPS mapping of the study site was made using Google earth pro software. Data analysis was
done by using SPSS 17.0 and XLSTAT 2009 softwares. MS- EXCEL 2007 was used for
data loading and graph preparation purpose.
RESULT
Total 19 different spp of butterflies of 4 different families have been recorded as regular
visitors during this study period on a single floral species Lantana camara (Table 1).
Maximum number of butterflies were recorded under the family Nymphalidae (9 species)
followed by Papilionidae (5 species), Pieridae (4 species) and Lycaenidae (1 species).
Dominance hierarchy is established on the basis of relative visiting frequencies (RVF) also
known as “encounters” on Lantana patches in the disturbed habitat (Table 2).
FAMILY Species Id Common Name CODES
1. Papillionidae Graphium
agamemnon (Linnaeus,
1758)
Tailed Jay TJ
2. Papillionidae Graphium doson (C. &
R. Felder, 1864)
Common Jay CJ
3. Papillionidae Papilio
demoleus (Linnaeus,
1758)
Lime L
4. Papillionidae Papilio
polytes Linnaeus, 1758
Common Mormon CM
5. Papillionidae Suastus
gremius (Fabricius,
1798)
Indian Palm Bob IPB
6. Nymphalidae Euploea core (Cramer,
1780)
Common Crow CC
7. Nymphalidae Ariadne
merione (Cramer,
1777)
Common Castor CCas
8. Nymphalidae Danaus Plain Tiger PT

10. chrysippus (Linnaeus,
1758)
9. Nymphalidae Vanessa
cardui (Linnaeus,
1758)
Painted Lady PL
10.Nymphalidae Tirumala
limniace (Cramer,
1775)
Blue Tiger BT
11. Nymphalidae Hypolimnas
bolina (Linnaeus,
1758)
Oriental Great
Eggfly
OGE
12.Nymphalidae Acraea
terpsicore (Linnaeus,
1758)
Tawny Coster TC
13.Nymphalidae Junonia
atlites (Linnaeus, 1763)
Grey Pansy GP
14.Nymphalidae Junonia
almana (Linnaeus,
1758)
Peacock Pansy PP
15.Pieridae Catopsilia
pomona (Fabricius,
1775)
Common Emigrant CE
16.Pieridae Appias
albina (Boisduval,
1836)
Common Albatross CA
17.Pieridae Appias
olferna (Swinhoe,
1890)
Striped Albatross SA
18. Pieridae Delias Common Jezebel CJez

11. eucharis (Drury, 1773)
19. Lycaenidae Chilades
pandava (Horsfield,
1829)
Plains Cupid PCup
Table 1: Checklist of butterflies recorded on Lantana camara during the study period.
Total number of visiting frequency (encounter) is 2184 (N= 2184) from February- May 2019.
Dependence level of different butterflies under different families based on the total
encounters is shown in the result (table 3, figure 4). Non-parametric Kolmogorov- Smirnov
(K-S) test is done to check the normality of the collected data of 19 different butterflies and
their encounters in monthly basis (table 4). As the data is in normal distribution; parametric
tests are preferred for further research questions. To test the variation between butterfly visits
(encounter) in different times and in different months, Analysis of Variance (ANOVA) test is
preferred. For the time specific study, two different time zones are selected as, time zone I (9
AM – 11AM) and time zone II (11 AM- 1 PM). Test statistics show that, there is no variation
exists between different butterfly visits in different time zones; as the p- value > 0.05 (table
5). Total work is done from February to May, 2019. To test the variation between different
butterfly visits (encounters) in different months further ANOVA test is performed.
Significant variation (p- value<0.05) is observed between only three butterflies visits out of
nineteen butterflies; namely, Common albatross(CA), Peacock pansy(PP) and Plains Cupid
(Pcup) in four different months ( table 6).
Sp. Id RVF Rank Value
T JAY 0.28021 1
C JAY 0.13369 2
LIME 0.12040 3
C.EMIGRANT 0.08740 4
S ALBATROSS 0.05998 5
C MORMON 0.04340 6
P.TIGER 0.04212 7
C ALBATROSS 0.03892 8
C. CROW 0.03617 9
C. CASTOR 0.03068 10
O G EGGFLY 0.02335 11
I.PALM BOB 0.02152 12
T.COSTER 0.02106 13
G. PANSY 0.01694 14
B.TIGER 0.01557 15
P.LADY 0.00915 16
C.JEZABEL 0.00915 17
P.PANSY 0.00549 18
P CUPID 0.00457 19
Total 0.99977

12. Table 2: Relative visiting frequencies (RVF) of different butterflies in accordance with
rank values.
Fig 3: Dominance hierarchy of different butterflies on Lantana camara.
The month of February was considered as 1, followed by March (2), April (3) and May (4).
ANOVA test is statistically significant on 3 butterfly visits in these four different months;
hence, a Post hoc analysis is performed to test where those variations exist, i.e. in which
months. Duncan Multi Range test (DMRT) at confidence level (alpha) 0.05 showed that three
different results for three different butterflies (table 7). In case of common albatross (fam.
Pieridae) there is a significant level of variation of encounters exist between the month of
March and May, and February and March (4>2=3>1).The encounters of Common albatross
in the month of March and April are equal comparatively. Peacock pansy (fam.
Nymphalidae) shows no variation between the months of February and March, but differs in
the month of March, April and May (4>3>2=1).From the DMRT it is observed that, the visit
frequencies (encounters) of Plains cupid is higher in the month of May followed by March
and February. The encounters are in same range in the months of February and April
(4>2>1=3). Lantana blooms in synflorecence as thickets. In the study site within the study
period three types of floral colors are found; namely yellow, pink and white. Data is taken as
C1=yellow, C2= pink and C3= white. There is a combination of flower colors, either white –
pink or yellow-pink blooms on same species Lantana camara.
From the present study, butterflies prefer yellow coloured flowers more, rather than pink or
white coloured flowers. From 2184 visits, 77.86% visit was on yellow flowers, 15.23% visit
was on white flowers and 6.90% visit was on pink flowers (figure 5). Individualistic flower
preferences were shown in figure 6; common albatross, blue tiger, Indian palm bob, peacock
0
0.05
0.1
0.15
0.2
0.25
0.3
RVF
Species
Dominance Hierarchy of Butterflies on Lantana
Plant
RVF

13. pansy and plains cupid butterflies do not prefer pink flowers. White flowers were to some
extent preferable by some butterflies. But, all the studied butterflies were largely dependent
on yellow flowers. To clarify a research question, is there any variation exist between
different butterflies visits in different coloured flowers an ANOVA test was performed. From
the result of the test statistics it was seen that, “there is a variation exist between different
butterflies visits in different coloured flowers”; as the p-value<0.05 (table 8).
The morphometrical measurements of both butterfly proboscis and corolla length (of flower)
were done by using microscopes and centimeter scales. Total 45 flowers were collected (from
10 different florets) from field. A one-sample t test was done to check that, is there any
significant difference between mean corolla lengths of flowers. Corolla length was denoted as
TFL (total flower length) in test statistics. Result showed that, there was a significant
difference exists between corolla lengths of flowers (table 9).
Total 9 different species of butterflies were collected from the field, each having 5 samples
for proboscis morphometrical studies. Mean proboscis lengths were shown in table 10. For
comparing mean differences of proboscis lengths of different butterflies, again a paired
sample t test was carried out. From 26 different pairs of butterflies, only 12 pairs showed
significant difference between mean proboscis lengths (table 11).
Thirteen different herbs were identified as potential nectar plants of the studied butterflies
(table 12).
1.
Amaranthus
spinosa
2.
Hyptis suaveolens
3.
Anisomelis
indica
4.
Ipomoea sp.

14. 5.
Cleome viscosa
6.
Parthenium
hystetophorous
7.
Cynodon
dactylon
8.
Physalis minima
9.
Euphorbia hirta
10.
Physalis sp
11.
Riccinus communuis

15. 12.
Tridax procumbens
13.
Vitis trifolia
TABLE 12. Plant species identification of the study site (potential host plants of
different butterflies).

16. FIGURE 7: Field gallery.

18. Family Encounters % Dependence on Lantana
Papillionidae 1309 59.93%
Nymphalidae 438 20.05%
Pieridae 427 19.55%
Lycaenidae 10 0.04%
N=2184
TABLE 3: Dependence level of different butterflies belonging from different families on Lantana.
FIGURE 4: Pie chart shows the dependence level of 4 different families of
butterflies.

19. TABLE 4. Kolmogorov- Smirnov Test statistics.

20. FIGURE 5. Flower preferences of all studied butterflies.

21. FIGURE 6: Individualistic flower preferences of studied butterflies. (N=2184)
-100
0
100
200
300
400
500
600
No of Visits
SPECIES
Individualistic Flower Preference of Butterflies
C 1
C 2
C 3

22. Sp
Id
TJ CE L CJ PT CC TC CA C
M
BT SA Cca
s
OG
E
IPB PL PP GP CJe
z
Pcu
p
df 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7
F -
valu
e
1.44
4
1.48
9
.74
0
1.33
7
1.08
7
1.08
8
2.60
4
.07
6
.55
8
.96
2
.20
3
.951 .993 1.12
9
.07
7
.33
3
1.24
8
.899 0.00
Sig .275 .268 .42
3
.291 .337 .337 .158 .79
2
.48
3
.36
5
.66
8
.367 .358 .329 .79
1
.58
5
.307 .380 1.00
0
TABLE 5: Time based ANOVA result.
Sp Id CA PP Pcup
df 7 7 7
F- value 18.24 11.33 7.167
Sig 0.008 0.020 0.044
P- value P<.05 P<.05 P<.05
TABLE 6: Month based ANOVA result.

23. TABLE 7. Duncan test result.
TABLE 8: Flower colour specific ANOVA test result.
Sp Id
Confidence level (α ) Duncan Test Statistics
CA 0.05 4>2=3>1
PP 0.05 4>3>2=1
Pcup 0.05 4>2>1=3

24. TABLE 9. One sample t test of corolla length.
Sp Id CE PT CA GP PP TC CJ BT SA
MEAN 1.1 1.1 1.1 1.1 1.1 0.9 1.0 1.1 0.9
TABLE 10. Mean proboscis lengths (cm.) of different butterflies.

25. TABLE 11. Paired sample t test showing differences of different butterfly proboscis lengths.

26. DISCUSSION
Variation of encounters of different butterflies on same nectar plant between two time zones
were not statistically significant but statistically significant level of variation do exist between
and within different months in butterfly encounters. Varying proboscis lengths were a very
good indicator for nectar consumption of different butterflies from a single plant.
Lantana camara is a small perennial shrub which can grow to around 2 m tall and form dense
thickets in a variety of environments (Sharma et al. 2008). L. camara has small tubular
shaped flowers, which each have four petals and are arranged in clusters in terminal
areas stems. Flowers come in many different colours, including red, yellow, white, pink and
orange, which differ depending on location in inflorescences, age, and maturity (Ram and
Mathur, 1984). Inflorescences are produced in pairs in leaf axils. In some forms, a yellow-
ring is present around the opening of the corollary tube (Sinha & Sharma 1984). A visual clue
to pollinating insects is provided by the yellow coloration of the flowers and colour change in
the flower might be stimulated after pollination (Mohan Ram & Mathur 1984). Lantana use
this strategy to propagate by offering some nectars to the butterflies.
Lantana is a major source of nectar for many species of butterflies and moths, which enhance
the pollination success of the species. Differences in corolla length, inflorescence diameter,
and number of flowers per inflorescenc attract some butterfly species to visit Lantana more
frequently than others. The present study clearly shows that the butterflies are comparatively
more dependent on yellow flowers (figure 6, table 8). Lantana camara has tremendous
adaptive measure and ability to propagate quickly as assisted by the creation of varieties
floral structures particularly of its coloration along with the distribution of nectaries in the
flowers thereby inviting large number of animal species, insects in particular to utilize the
resources produced by Lantana for their own sustenance. It is true that, Lantana has earned
the tag of a weed mostly by its invasive nature and destruction of plant diversity in the
economic region of forest or in degraded and waste lands. However, Lantana has established
itself in the ecosystem and is functioning as one of its components in the food web thereby
establishing mutual interdependence.
CONCLUSION
From the present study it was revealed that, there are nineteen different types of butterflies
dependent on Lantana camara in the study area, within these months. The dominance of
Papilionidae butterflies were seen from the results. Important to note that, Nymphalidae
butterflies share a very good number in the community, followed by Papilionidae, Pieridae
and Lycaenidae butterflies. All these butterflies show a greater preference on yellow coloured
flowers, followed by white and pink flowers on a single floret. So many potential nectar
plants were present, but they choose Lantana mostly. The biochemical aspects of the nectar
sugar concentration and the nectar quality analysis of the Lantana plant is a thirst area for
further research purpose. The present work also shows that, the proboscis lengths of the
butterfly vary within and between same and different families. Four months dissertation work
gave a basic understanding, but not a concluding scientific statement on this particular plant
animal interaction.

27. ACKNOWLEDGEMENT
I am indebted to the Department of Zoology, West Bengal State University for giving me the
opportunity to take field biology as my optional elective. I owe my vision and
accomplishment of the dissertation work on butterflies and Lantana camara to my supervisor
Prof. Narayan Ghorai, whose valuable guidance, enormous liberty to do the field work
regarding the project planning and continuous encouragement and support throughout my
study has made this work a fruitful reality. My heartiest gratitude goes to some names like
Shawon Bayen, Bedanta Das, Ankita Ghosh, Aritra Bhattachrya, Kuladeep Roy and Arunava
Mukherjee for their kind cooperation in various aspects. Last but not the least, my sincere
thanks goes to my family for the successful fulfilment of the project work.
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